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JOURNAL OF GENETICS

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JOURNAL OF GENETICS

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DIRECTOR OF THE JOHN IN'NES HORTICULTURAL INSTITUTION
AND

R. C. PUNNETT, M.A., F.R.S.

ARTHUR BALFOUR PROFESSOR OF GENETICS IX THE UNIVERSITY OF CAMBRIDGE

Volume III. 1913— 1914

Cambridge :
at the University Press

.1914

_->

(Tiimbntigr :

rRINTED BY JOHN CLAY, M.A.
AT THE UNIVERSITY PRESS

CONTENTS.

No. 1 (June, 1913)

PAOE

L. DoNCASTER. On an Inherited Tendency to produce purely Female
Families in Abraxas grossulariata, and its Relation to an Ab-
normal Chromosome Number ....... 1

L. DoNCASTER. On Sex-limited Inheritance in Cats, and its bearing

on the 8ex-limited Transmission of certain Human Abnormalities 1 1

H. H. New.man. Five Generations of Congenital Stationary Night-

Blindness in an American Family. (With 2 Text-Figures) . 25

Geoffrey Smith and Mrs Haig Thomas. On Sterile and Hybrid

Pheasants. (With Plate I) 39

N. Barlow. Preliminary Note on Heterostylism in Oxalis and

Lythrum. (With 1 Text-Figure) .53

Philippe de Vilmoris. Sur une Race de Ble Nain Infixable.

(With Plate II, and 8 Text-Figures) 67

No. 2 (September, 1913)

R . C. PuNNETT. Reduplication Series in Sweet Peas ... 77
Caroline Pellew. Note on Gametic Reduplication in Pi»um . 105
J. C. F. Fkyer. Preliminary Note on some Experiments with a

Polymorphic Plasmid. (With Plate III) 107

Edward N. Wentworth. The Segregation of Fecundity Factors in

Drosophila . . . . . . . . . .113

J. A. Jenkins. Mendelian Sex-Factors in Man . . . .121

Charles Todd. On the Recognition of the Individual by Haemolytic

Methods . . 123

C. J. Bond. Some Points of Genetic Interest in Regeneration of

the Testis after Experimental Orchectomy in Birds. (With

Plates IV and V) 131

vi Contents

No. 3 (February, 1914)

PAGE

Robert K. Nabours. Studies of Inlieritance and Evolution in Oitho-

ptera I. (With Plate VI, 3 Text-Figures, and 2 Diagrams) . 141

C. W. Richardson. A Preliminary Note on the Genetics of Fragaria.

(With Plate VII and 4 Text-Figure.s) 171

W. E. Agar. Parthenogenetic and Sexual Reproduction in ISimo-

CPi>halus vetiilus and other Cladocera . . . . .179

E. S. Salmon. On the Appearance of Sterile "Dwarfs" in Ifumulus

Lupulus L. (With Plates VIII and IX) . . . .195

I. B. J. SoLLAS. Note on the Offspring of a Dwarf-bearing Strain

of Guinea Pigs .......... 201

C. J. Bond. On a Case of Unilateral Development of Secondary
Alale Characters in a Pheasant, with Remarks on the Influence
of Hormones in the Production of Secondary Sex Characters
(With Plates X— XIII) 20.5

H. Drinkwater. Minor-Brachydactyly. No. 2. (With Plates

XIV— XVI and 1 Chart) 217

P. G. Bailey. Primary and Secondary Reduplication Series. (With

1 Text-Figure) 221

No. 4 (April, 1914)

J. W. H. Harrison and L. Donca.ster. On Hybrids between Moths
of the Geonietrid Sub-Family Blatoninae, with an Account of
the Behaviour of the Chromosomes in Gametogenesis in Lycia
(Bis/on) Hirinrin, Itlnjsia (^Nytisla) Zoiiar'm and in their Hybrids.
(With Plates XVII and XVIII) 229

Onera a. Merritt Hawkes. On the Relative Lengths of the Fir.st
and Second Toes of the Human Foot, from the Point of View of
Occurrence, Anatomy and Heredity. (With Plates XIX — XXI
and 6 Text-Figures) 249

Rose Haig Thomas. The Transmission of Secondary Sexual
Characters in Pheasants (Witli Plates XXII— XXVI and
2 Text-Figures) 275

W. Bow.\TER. Heredity of Melanism in Lepidoptera. (With Plate

XXVII) . . ■ 299

Vol. 3, No. 1

June, 1913

JOURNAL OF GENETICS

EDITED BY

W. BATESON, M.A., F.R.S.

DIRECTOR OF THE JOHN INNES HORTICULTURAL INSTITUTION
AND

R. C. PUNNETT, M.A., F.R.S.

ARTHUR BALFOUR PROFESSOR OF GENETICS IN THE UNIVERSITY OF CAMBRIDGE

CAMBRIDGE UNIVERSITY PRESS

C. F. CLAY, Manager

LONDON : FETTER LANE, E.C.

EDINBURGH: lOO, PRINCES STREET

also

H. K. LEWIS, Ij6, GOWER STREET, LONDON, W.C.

WILLIAM WESLEY AND SON, 28, ESSEX STREET, LONDON, W.C.

PARIS ! LIBRAIRIE HACHETTE & c"^.

BERLIN : A. ASHER & CO. LEIPSIC : BROCKHAUS

CHICAGO : THE UNIVERSITY OF CHICAGO PRESS
BOMBAY AND CALCUTTA : MACMILLAN & CO., LTD.

Price Ten Shillings net

[Issued June 25]

The University of Chicago Press

Heredity and Eugenics. By John M. Coulter, William E.
•Castle, Edward M. East, William L. Tower, and Charles B.
Davenport. 312 pages, 8vo, cloth; 10s. net.

Five leading investigators, representing the University of Chicago, Harvard
University, and the Carnegie Institution of Washington, have contributed to this
work, in which great care has been taken by each contributor to make clear to the
general reader the present position of evolution, experimental results in heredity in
connection with both plants and animals, the enormous value of the practical
application of these laws in breeding, and human eugenics. Technicalities of
language have been avoided, and the result is an instructive and illuminating
presentation of the subject for readers untrained in biology as well as for students.

Contents: I. Kecent Developments in Heredity and Evolution: Generallntroduetion.

II. The Physical Basis of Heredity and Evolution from the Cytological Standpoint (John
Merle Coulter, Professor and Head of the Department of Botany, the University of Chicago).
HI. The Method of Evolution. IV. Heredity and Sex (William Ernest Castle, Professor
of Zoology, Harvard University). V. Inheritance in Higher Plants. VI. The Application
of Biological Principles to Plant Breeding (Edward Murray East, Assistant Professor of
Experimental Plant Morphology, Harvard University). VII. Recent Advances and the
Present State of Knowledge concerning the Modification of the Germinal Constitution of
Organisms by Experimental Processes (William Lawrence Tower, Associate Professor of
Zoology, the University of Chicago). VIII. The Inheritance of Physical and Mental
Traits of Man and their Application to Eugenics. IX. The Geography of Man in
Relation to Eugenics (Charles Benedict Davenport, Station for Experimental Evolution,
Carnegie Institution of Washington).

British Medical Journal. Those who are desirous of arriving at an estimate of the
present state of knowledge in all that concerns the science of genetics, the nature of
the experimental work now being done in its various departments,... and the prospects,
immediate or remote, of important practical applications, cannot do better than study
"Heredity and Eugenics."

The Nation, New York. "Heredity and Eugenics" may be heartily recommended to
readers seeking, as beginners, to get in touch with the discussion of these subjects. ...In
most of the lectures there is an admirable reserve, not to say skepticism, in the treatment
of large questfons which the public is often misled to regard as already and finally settled.

The Mechanistic Conception of Life. Biological Essays. By

Jacques Loeb, Head of the Department of Experimental Biology,

Rockefeller Institute for Medical Research. 238 pages, 12mo,

cloth ; 6s. net.

Professor Loeb's experimental researches at the University of Chicago, the
University of California, and the Rockefeller Institute for Medical Research, ensure
this new collection of his latest conclusions a wide reading. The author's purpose
in this book is to discuss the question whether present knowledge gives any hope
that hfe may be unequivocally explained in physico-chemical terms. An affirmative
answer, he thinks, will necessitate a reconstruction of our social and ethical life on
a scientific basis. This volume is a popular presentation of the results of the
author's investigation.s, including his successful experiments in chemical fertUizatiou.
The wide range of his discussion is seen in the following list of contents : I. The
Mechanistic Conception of Life. II. The Significance of Tropisms for Psychology.

III. Some Fundamental Facts and Conceptions concerning the Comparative
Physiology of the Central Nervous System. IV. Pattern Adaptation of Fishes
and the Slechanism of Vision. V. On Some Facts and Principles of Physiological
Morphology. VI. On the Nature of the Process of Fertilization. VII. On the
Nature of Formative Stimulation (Artificial Parthenogenesis). VIII. The Prevention
of the Death of the Egg through the Act of Fertihzation. IX. The Role of Salts
in the Preservation of Life. X. Experimental Study of the Influence of Environment
on Animals.

The Cambridge University Press

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Volume III JUNE, 1913 No. 1

MRY

ON AN INHERITED TENDENCY TO PEODUCE eoTlJlcAL
PURELY FEMALE FAMILIES IN ABRAXAS (i^Kum.^
GROSSULARIATA, AND ITS RELATION TO
AN ABNORMAL CHROMOSOME NUMBER.

By L. DONC aster, M.A.,

Fellow of Kiiufs College, Cambridge.

I. The Inheritance of Unisexual families.

It is well-known that in Lepidoptera, families are occasionally
produced which are all or nearly all of one sex. This has sometimes
been observed as the result of hybridizing distinct species (e.g. Tutt,
Trans. Entom. Soc. 1898, p. 17. Three matings of Tephrosia histortata $
X T. crepuscularia ^ gave 40 j/, 0 ?; 58 (/, 0 ? ; 60 ^, 1 ?), but not
rarely families consisting wholly of males or of females have been
recorded when both parents have been apparently normal members of
the same species (cf Lamborn's experiments with Acraea encedon
(Proc. Entom. Soc. 1911, p. liv) referred to below).

In 1908 I received fi-om the Rev. G. H. Raynor a batch of larvae of
Abraxas grossulariata, obtained by mating a wild female with a lacti-
color male (No. '07.19^). From these I reared 19 moths, all lacticolor
females, and heard from Mr Raynor that from the same family he had
bred 21 lacticolor females, making 40 in all. Of my females I paired
four, one with a lacticolor male, three with grossulariata males all of
which were heterozygous for lacticolor. The pairing with a lacticolor
male (^08. 2) gave 27 lacticolor females and no males ; from one of the
other pairings no images were reared ; the remaining two gave respec-
tively 40 c/", 23 ? , and 5 c/, 2 ? . Of the 27 females in family '08. 3 I
paired six, three with lacticolor males, three with grossulariata males.

' The reference numbers of purely female families are italicised.
Journ. of Gen. in 1

2 Female Families in Abraxas-

Of these males, one (mating '09. 12) proved to be homozygous for gros-
sulariata, a second heterozygous ('09. 7), and from the third no imagos
were reared. From the two successful matings with grossulariata
males, the results were 52 ^, 25 $ and 42 </, 25 ?. The matings with
lacticolor mviXes were '09. 8, which yielded 11 (/,15 $; '09. 15, one female
only; and '00.18 with 11 females. The single female from '09. 15 was
paired with a lacticolor male from '09. 7, but laid no eggs ; of the
eleven females from 'Of). IS ail were paired, but three produced hardly
any eggs and no larvae (two of the males used in these infertile matings
were lacticolor- from '09. H, the thinl a grossulariata frcjm '09. 12, i.e. all
were sons of females of the unisexual family 'OS. J). Of the eight
fertile pairings, two (^10.21, '10.24) were with lacticolor males; the
male of '10. 31 was unrelated and from 80 eggs 24 females were reared,
with no males; the male of '10. 24 was from '09.7 (.son of '08.2) and
from 100 eggs were reared 22 </, 25 $. Of the six fertile pairings
with grossulariata males, one ('10. 19) was with a wild male and gave
from about 75 eggs 25 {/", 20 ^ ; two were with homozygous gross.
(unrelated) males and gave from 56 and 80 eggs 21 (Z", 23 % and 4^^ , 5 ^
('10. 25 and '10. 27). The other three were with heterozygous gross.
males; in one ('10. 15), in which the male was unrelated, 32 jf and 27 $
were produced; the other two, '10.22 and '10.28, in which the male in
each case was from '09. 12 (sons of '08. 2), produced from 19 and 91 eggs,
4 $ (3 gross. 1 lact.) and 62 % (34 gross. 28 lact.). In these last matings,
purely female families including grossulariata as well as lacticolor were
produced for the first time.

Another important point arose from the 1910 matings. A lact.
female of '09. 8 (a bisexual family, whose mother belonged to the
unisexual family '08.2) was paired with an unrelated lact. </, and
produced 69 ?, 4c/' ('10.17), and in another family ('10. 10) descended
on both sides more or less nearly from unisexual fiimilies (see Table of
Matings) 14 $ and 2;/' were produced. The 1910 pairings showed
therefore (1) that there is apparently some tendency to infertility
among females of unisexual families, at least when mated with related
males; (2) that unisexual families including grossidnriata individuals
can be produced ; and (3) families with enormous preponderance of
females were reared from parents which belonged to normal broods
themselves, when one or both were descended from unisexual families.

It is not necessary to describe in detail the matings made in 1911 ;
the results of those which are important for the present purpose are
given in the table. It will be seen that of twelve matings with females

L. DONCASTER 3

from the unisexual family '10. 28, three (7i. i-5, '11. 10, '11. 36) gave only
female offspring, and that of these families one was by a wild male, the
other two by males unrelated to the female. Half the matings of
females from '10.38 were nearly or totally infertile, in two cases ('11.27
and '11. 29) when the male was unrelated to the female. Similar
infertility was found in two of the three matings of females from '10. 31,
and in four matings (not included in' the table) of females from '10. 10
and '10. 17 (families in which there was a great preponderance of
females).

A fresh point of importance also appeared. In three matings {'11. 35,
'11. 9 and '11. 4) of which the female parent in each case was daughter
of a female from '09. 18, and the male in two cases a son of a female
from '00. 18, and in the third a member of the preponderatingly female
family '10. 17, only females were produced, although both parents were
themselves members of bisexual femilies.

It is clear, therefore, not only that the tendency to produce purely
female families may be transmitted direct from mother to daughter,
even when the male parent is unrelated, but also that females of
bisexual families directly descended from unisexual families may produce
only female offspring, at least when mated with males of similar origin.
Further, of the females of unisexual families, about half have only
female offspring, and the remainder have offspring of both sexes.

Several ^Joints of importance remain to be determined, and it is to
be hoped that the results of the pairings made in 1912 will clear them
uj). The more important are (1) whether females of bisexual families,
whose mothers belong to unisexual families, can have only female
offspring when mated with unrelated males ; (2) how the families with
great preponderance of females are related to the completely unisexual
broods ; and (3) whether mortality in the egg or larval stages has any
relation to the production of only female offspring. Experiments to
test all these points are in progress. It was hoped that the third
would be settled by the 1911 matings, but the exceptionally hot dry
weather at the time of hatching caused great mortality in the very
early stages. Although in some cases a large proportion of the eggs
failed to develop, in other batches from which only females were pro-
duced almost every egg hatched, and the case of '10. 38, in which 62
females were reared from 91 eggs, strongly suggests that the production
of only female offspring is not due to the dying off of the male larvae.
Further evidence in the same direction is provided by the preliminary
account of the chromosomes in unisexual broods which follows. Another

1-2

Female Families in Abraxas

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6 Female Families in Abraxas

point of interest should be noticed — that in the unisexual families the
ordinary rides of sex-limited transmission are followed. A grossulariata
female, mated with a lacticolor male, normally produces lacticolor female
and f/7-ossidanata male offspring; if the fiimily consists entirely of
females, these ai"e all lacticolor, and the grossulariata character is not
transmitted at all (cf. ftxmilies '07. 19, '11. 9 and '11. 4).

It has been mentioned that a considerable proportion of matings of
females from unisexual families, especially when the male is related, are
infertile. In 1912 I dissected most of the females paired, after the
eggs were laid, in order to see whether the ovaries were in any case
abnormal. Although in three cases I have found ovaries containing no
oocytes in larvae dissected for histological examination, I found no
abnormal ovaries, with the exception of one which had six egg-tubes
instead of four, in the moths which had been paired. In several cases,
however, in which the eggs were infertile I found that the spermatheca
was nearly or quite empty, showing that copulation had not been
successful, but this cannot be the .sole cause of the infertility, for in
other cases in which the eggs foiled to develop the spermatheca was
well filled.

Before passing on to the account of the cytology of the unisexual
families, it should be mentioned that my case of a heieditary tendency
to produce families consisting of females only ajjpears to be closely
comparable with that discovered by Dr Lamborn in Acraea encedon, of
which a preliminary account has been published (Proc. Entoni. Soc.
1911, p. liv). I hear fi'om Prof Poulton that so far as the experiments
have gone at present, females from purely female families always have
only female offspring ; those from bisexual families have both males
and females. If this is confirmed, it will show that the two cases are
not exactly similar.

II. The Chromosomes in Unisexual families.

In recent papers' I have shown that the unreduced number of
chromosomes in both male and female of Ab7-axas grossulariata is
normally 56, and that there is no recognisable difference between the
type and va7\ lacticolor. In determining the chromosome number in
the oogonia of certain lacticolor females, however, I found several
perfectly clear figures in which only 5H could be counted, although in

' Joiiiwil (ifGe}ielics, i. 1911, p. 179, and ii. 1912, p. 189.

L. DONCASTER 7

other lacticolor females there were 56. The females in which I first
found 55 belonged to a family descended from unisexual stock, and it
occurred to me that possibly the absence of one chromosome might be
correlated with the tendency to produce only female offspring. In the
autumn of 1912, therefore, I dissected all the larvae which attained a
sufficient size, and preserved the ovaries and testes'. The sexes are
quite easily i-ecognisable on dissection when the larvae are less than
half-grown, and the oogonial and spermatogonial divisions are more
numerous and clearer at that stage than later. In male larvae sperma-
tocyte divisions occur from about half-grown larvae onwards. The
methods used were the same as those described in my former papers ;
in counting chromosomes every figure has been dravm twice, at
intervals of a day or usually more, in order to get independent con-
firmation of each count.

Of the femilies obtained from fenmles of unisexual broods only two
provided any considerable number of larvae which grew to a sufficient
size, '12. 8, from a gross. $ ex. '11. JG x gross. ^ ex. 11. 11, and '12. 25,
from gross. J ex. '11. lUxlact. ^ ex. '11. 14. From '12. 8, 83 larvae were
dissected, every one a female ; from '12. 25, 14 were dissected, 10 f/", 4 $ .
Several other families yielded from two to six larvae, some of which
will be referred to below. Since in normal families the larvae which
feed up in the autumn are much more often males than females, it may
be taken as highly probable that '12. 8 is a unisexual family ; where
only six or fewer females were dissected, it is possible that some males
will ajjpear later.

Of the 33 larvae (all females) of '12. 8 which were dissected and
their ovaries fixed, only 14 provided figures of oogonial divisions which
could be counted with any accuracy. Of these, nine provided altogether
eighteen figures in which 55 chromosomes could be counted with almost
complete confidence, and of these two or three especially are so perfectl}^
clear that no doubt about the number is possible. In a number of
other figures in these ovaries 55 is the most probable number, but in
counting chromosomes, when their number is large, a few really trust-
worthy figures are worth much more than many rather doubtful ones.
In addition, out of the fourteen larvae which yielded good or fair
figures, two showed four figures in which only 54 can be seen (none of
them first-class figures), and three showed four figures in which no

' The majority of yrossulariata larvae cease feeding when quite small in the late
summer, and hibernate at that stage. A small and varying number feed up and pupate
in the autumn. All the latter were dissected, usually when about half-grown.

8 Female Families in Abraxas

decision between the numbers 55 or 56 was possible. In one of the
latter, however, several figures gave 55 and only one 56, and from a
slight difference in staining there is reason for believing that one of the
bodies counted as a chromosome among the 56 is really an extra-
nuclear body at the edge of the chromosome group. Among the
offspring of pairing '12. S, therefore, we have over 20 figures in which
the number 55 is certainly or almost certainly correct, four in which
only 54 can be seen, and none in which there is conclusive evidence of
as many as 56. The number 55 is also the most probable in several of
the ovaries in which the figures are not sufficiently clear to be used as
evidence.

Two other families, '12. 1 and '12. 32, provided sufficient larvae to
give considerable jjrobability that they will turn out exclusively female.
Both are from females belonging to unisexual families; '12.1 from
lact. % ex. '11. 4. X lact. </ ex. '11. 6, and '12. 32 from htct. % ex. '11. 9
X wild (/'. Of '12. 1, four larvae were dissected, all females; two pro-
vided ovaries with three countable figures, all with 55. In '12.32, .six
larvae were dissected, again all females ; four of these show good or fair
figures, two figures with 55 clearly, and four others with 55 probably.
In the three families, therefore, there are about thirty good equatorial
plates showing 55 chromosomes, and only eight in which 55 or 56 might
be counted with equal probability.

In only one other family the mother of which belonged to a
unisexual brood have any definite results been obtained up to the
present time. Family '12. 25, from gross. % ex. '11. IGxlact. </ ex. '11.14,
has given ten male and four female larvae. One of the females provides
an equatorial plate with 56 chromosomes shown quite clearly ; two
others probably have 56 and one probably 55 but possibly 56. In the
case, therefore, of bisexual families of which the mother belongs to a
unisexual family it is clear that some at least of the females have 56
chromosomes, but it is probable that others have 55.

The testes of a number of males ti'om families of which the mother
belonged to a unisexual family have been examined, chiefiy from '12.25
and '12. 21 B (the mother of the latter was gross. $ ex. '11. 36, the father
lact. cf ex. '11. 24 A). The chromosomes in spermatogonial divisions are
less easy to count than in the oogonial, but most of the really clear
cases have given 56, and both primary and secondary spermatocytes
have 28. As far as the work has gone at present, therefore, there is no
sufficient reason for regarding the males in bisexual families directly
descended from unLsexual families as differing in their chromosome

L. DONCASTER 9

number from the normal. It should be mentioned that in the males
of '12.25, almost constantly some of the spermatocytes are binucleate
or more rarely have four nuclei, and these give rise to binucleate or
quadrinucleate spermatozoa. I have not traced the stage at which this
condition arises.

It would be premature to discuss at this early stage the meaning of
the observations described above. It appears probable that all the
females of a unisexual family have .5.5 chromosomes instead of 56.
Those which have only female ottspring must therefore transmit only 27
to each of their offspring, and one chromosome is thus not accounted
for. An attempt will be made when the moths hatch to find out
whether one chromosome is eliminated at the maturation of the egg,
and until that point is decided it is useless to speculate on the relation
of the chromosome number to sex-determination.

There is one point of considerable theoretical importance, however,
which should be referred to. A gi-ossulariata female which has only
female offspring does not transmit the gross, factor to any of them — the
mother receives it fi-om her male paient, but does not transmit it to her
daughters, and since in this case she has no sons, she does not transmit
it at all. In some way, therefore, it is eliminated, perhaps most
probably at the maturation of the egg. She also receives 28 chromo-
somes from her male parent, but transmits only 27 to her offsf)ring ;
one chromosome must therefore be eliminated. It seems, therefore,
highly probable that this chromosome, which is not transmitted by the
female whose offspring are all females, may be the determin(5r or bearer
of the grossuhiriata factor. It is tempting also to suggest that this
chromosome is a sex-determiner — that if it is receive<l from the mother
and a corresponding one from the father, the zygote becomes a male,
and if it is received from the father only, it becomes a female. A diffi-
culty arLses, however, from the fact that some females at least, whose
mothers belonged to unisexual families, have 56 chromosomes, as has
been shown in the case of family '12. 25. It is probably best, therefore,
to reserve further consideration of this part of the subject until our
knowledge of the facts is more complete.

Summary.

1. Families consisting wholly of females have appeared in six
successive generations of Abraxas grossidariata ; they may consist of
var. lacticolor, of grossvhmata, or of equal numbers of each, according
to the constitution of the male parent.

10 Female Families^ in Abraxas

2. In some cases unisexual families have been produced when both
pai-ents belonged to bisexual families descended directly from unisexual
families. In other cases such matings have given families with gi-eat
preponderance of females, but some males.

3. About half the females of a unisexual family have only female
offspring. Infertility in such families is frequent.

4. The normal number of chromosomes in both sexes, whether
gross tdariuta or lacticolor, is 56. Females of unisexual families, how-
ever, have often, and probably always, only 55.

5. Females of bisexual familie.s, descended directly from unisexual
families, have in some cases 56, probably in others 55 chromosomes.
As far as is known, males always have 56.

6. It is suggested that the elimination of one chromosome may be
correlated with the elimination of the grossuluriata factor. The
investigation is not yet far enough advanced to give definite evidence
of the relation between the behaviour of the chromosomes and the
phenomena of sex-determination in this species.

ON SEX-LIMITED INHERITANCE IN CATS, AND ITS
BEARING ON THE SEX-LIMITED TRANSMIS-
SION OF CERTAIN HUMAN ABNORMALITIES.

By L. DONCASTER, M.A.,

Fellow of Kinff's College, Cambridfje.

Sex-limited inheritance is now kno\\Ti to be a widespread pheno-
menon, but several problems of importance in connexion with it still
remain obscure. It has been shown to exist in Lepidoptera and Diptera,
in Birds and Mammals, and falls into two classes, according to whether
the sex-limited transmission is by the male or the female. In Abraxas
(Lepidoptera) and in the Fowl, Canary and Pigeon the female is found
to be normally heterozygous for certain characters, and with rare
exceptions transmits them to her male offspring only ; in Drosophila
(Diptera) and in Man and the Cat the male is similarly heterozygous
and transmits certain characters chiefly or only to his daughters. The
simple conclusion from these facts would be that in Lepidoptera and
Birds the sex of the offspring is determined by the egg, and in Diptera
and Mammals bj' the .spermatozoon. There is evidence, however, that
the sex-ratio in Drosophila is partly at least dependent on the female
parent^ and evidence pointing in the same direction has been brought
forward in the case of Man. It is plain therefore that a full knowledge of
sex-limited inheritance is of fundamental importance for the solution of
the problem of the determination of sex. It is also of great importance
from a somewhat different point of view. The hypothesis has been
widely accejDted that not only is sex determined by the so-called sex-
chromosomes, which have been chiefly studied in Insects, and are known
to exist in many other groups of animals, but also that the factors for
sex-limited characters are also borne by these .sex-chromosomes. Simple

' Moenkhaus, W. J., .Tourn. Morphol. Vol. xxii. 1911, p. 23; Eawls, Biol. Bulletin,
XXIV. 1913, p. 11.5. [Morgan, Science, Vol. xxxvi. p. 718, Nov. 22, 1912, has given an
explanation of Miss Rawls's results which is not inconsistent with sex determination by
the spermatozoon in Drosophila. Note added March 5, 1913.]

12 Sex-Limited Inheritance in Cats

and attractive as this hypothesis is, it cannot be regarded as established
until certain contradictory facts have been explained. Apart fioni the
fact that an odd chromosome has been described by Gnyer in the
spermatogenesis of the Fowl', although in this case it is the female and
not the male which shows sex-limited transmission, the hypothesis has
to fiice two chief difficulties. The first and less important is that no
case is certainly known in which an odd or unequally paiied chi-omosome
exists in the female'-. This difficulty is not serious, since it is (juite
possible that a physiological difference niay exist between the members
of a pair of chromosomes, even though it may not be made visible by
our rather crude cytological methods. The second difficulty is more
fundamental, and arises from the fact that in the Canary and Pigeon at
least, and possibly in other cases, the sex-limitation of the transmission
of the characters studied is not absolute. A dark-eyed female Canary
does not transmit the factor for tln' dark eye only to her sons; for when
mated to a red-eyed cock a small propurtion of the female offspring are
also dark-eyed'. In the case of sex-limited transmission by the male
no certain evidence of exceptions of this kind has been published,
although some of Morgan's exceptional cases in Drosopliila are most
easily explained on the assumption of a failure of the normal sex-limited
transmission*.

The position is therefore this. In one order of Insects and one class
of Vertebrates it appears that sex is determined by the egg; in a
second order of Insects and a second class of Vertebrates the same kind
of evidence suggests that sex must be determined by the spermatozoon.
In the second group the evidence for sex-determination by the sperma-
tozoon is supported by the visible existence of unpaired sex-chromosomes,
giving rise to two kinds of spermatozoa, and yet in buth Drosopliila and
Man thei-e are indications that sex-determination is dependent on or
influenced by the mother. Secondly, it appears that in some cases at
least sex-limitation in the Iransniission of characters is not absolute,
but is partial as in the case of gametic coupling of other characters,

' Anat. Anzeiger, Vol. xxxiv. p. 573.

- Baltzer's description of such dimorphism amonp; the eggs of Echiiioids can hardly
be regarded as proved, until tlie discrepancies between his results and those of Tennent
have been removed. [J. Seiler, Zool. Anz. xli. p. 24G, has just published a preliminary
note indicating possible chromosome-dimorphism in the eggs of the Moth Phraijinatvliia.
Mar. 1913.]

'■> Durham and Marryat, Reports to Evolution Committee, Ilniinl Soc. iv. 1908, p. 57.

* E.g. the occurrence of a miniature y in I<\ from rud.-min. { ■: long male. Zeitschr.
f. i/tdult. Abslamm. vii. 1912, p. 333.

L. DONCASTER 13

and yet the hypothesis most widely adopted, especially in America, for
the explanation of sex-limited transmission assumes that the sex-factor
and the factor for the sex-limited character are borne by the same
chromosome.

In the hope of obtaining fresh evidence on these two points I have
collected and analysed a number of pedigrees of sex-limited affections
in Man — Colour-blindness, Night-blindness, Nystagmus and Haemo-
philia'. A preliminary examination of these showed that all four
affections had the same characteristics as regards their inheritance —
an apparent disturbance of the sex-ratio among the offspring of
transmitting females, an excess of affected over unaffected males in
affected fraternities, and occasional exceptions to the ordinary rule of
sex-limited transmission among the children of affected males. On
tabulating a considerable series of jjedigrees, however, it became clear
that there were several imj)ortant sources of error, which it is difficult
to eliminate, arising partly from the incompleteness of the pedigrees,
and partly from the fact that it is impossible in most cases to know
that a woman is a transmitter of the affection unless she has at least
one affected son. While I was engaged on the work, I received a paper
by Lenz^ which points out that the apparent excess of males in affected
fraternities, and also the excess of affected over unaffected sons, is
possibly due to the fact that of necessity the totals of offspring of
transmitting females are compiled from fraternities including at least
one affected male. Fraternities consisting chiefly of daughters, with no
affected son, are excluded, since they provide no evidence that the
mother is a transmitter of the affection, and thus there arises a
preponderance not only of males over females, but of affected over
unaffected males. With the object of testing this suggestion I have
tabulated the fi'aternities from my data for the four affections in which
there are at least seven children, and find that in these totals the
apparent excess of males is much reduced in each case, and in some
vanishes altogether. The prepondei'ance of affected over unaffected

^ Colour-blindness — Nettleship, Tram. Ophthalm. Soc. xxvm. 1908, p. 220 ; xxvi.
1906 ; and unpublished cases kindly given me by Mr Nettleship and by Mr S. P. Hayes
of Mount Holyoke College, U.S.A.

Night-blindness and Nystagmus — Nettleship, Trans. Ophthalm. Soc. xxix. 1909, p. Ivii;
Jixxi. 1911, p. 1-59 ; xxvni. 1908, p. 220 ; xxxii. 1912, p. 21 ; and Royal London
Ophthalm. Hospital Reports, xvn. p. 333.

Haemophilia — Bullock and Fildes, Treasury of Human Inheritance, Parts v. and vi.
London, 1911.

- Lenz, Ubcr die Krankhaften Erbanlagen des Mannes. Jena (G. Fischer), 1912.

14 Sex-Limited Tnheritaace in Cats

also disappears in the ease of Colour-bliiidness and Night-blindness, but
remains ahnost unaltered in the Nystagmus totals. In the case of
Haemophilia it varies greatly ; in some families there is in each
fraternity a great excess of affecteds, in others eijuaJity or deficiency ;
the totals when oidy large fraternities are included show a smaller
excess than when all fraternities of evei'y size are added together. It
is the possible existence of this excess of affected individuals in the
offspring of transmitting females which is chiefly of importance for the
ptu'pose for which the enquiry was undertaken. For it has been pointed
out above that in some cases at least sex-limitation is not absolute, but
is partial like gametic coupling of other characters, and it seems at
least possible that the excess of affected over normal males among the
sons of transmitting females might be due to partial coupling of the
factor for the disease with a sex-factor, among the gametes of the female
parent. We know that the factor for the affection is absolutely or
almost absolutely coupled with a sex-factor in the gametes of the
affected male, for he transmits the factor only, or almost entirely, to his
daughters, his sons being very rarely if ever affected. If then we also
found that a woman bearing this fiictor transmitted it chiefly to her
sons, we should have absolute sex-limited transmission by one sex,
and partial sex-limited transmission by the other sex, in the same
species. A basis would thus be provided for a reconciliation of the two
types of sex-limited inheritance, exemplified respectively by Abraxas
and Birds on the one hand, and by Drosupldla and Mammals on the
other.

If this were the case, the high ratio of affected sons of a trans-
mitting woman should be associated with a low ratio of transmitting
to non-transmitting daughters. My material gives no evidence that
this is the case; although the data for determining whether the
daughters of a transmitting woman do or do not transmit are veiy
inadequate, they suggest that the transmitting daughters are more
numerous than they should be if the excess of affected over normal sons
were due to gametic coupling of the factor for the disease with a sex-
factor, in the gametes of the transmitting mother. The families which
have an exceptionally high ratio of affected males also have a high ratio
of transmitting daughters, instead of a lower ratio as would be expected
on the gametic coupling hypothesis. It is hardly possible, therefore,
with the data at present available, to come to any definite conclusion
about the nature of the transmission of these hmnan abnormalities. It
can only be said that there is in some pedigrees an excess of affected

L. DONCASTER 15

over unaffected males, but that this is not a constant feature, and when
it occurs it is improbable that it is duo to partial sex-limitation among
the gametes of the transmitting female. Further, it is probable, if not
certain, that occasional exceptions occur, suggesting that an affected
man may very rarely transmit the fiictor for the affection to a son, and,
correspondingly, may occasionally fiiil to transmit it to a daughter.

The difficulties in the way of an elucidation for the human cases
arise chiefly from the impossibility of distinguishing a transmitting
female except by the fact that she has affected sons. In the hope of
throwing further light on the question, I compared my summaries of
human pedigrees with data which I have collected during several years
of colour-inheritance in Cats, and find that they are in most respects so
closely similar that I believe the inheritance of certain characters in the
Cat may provide a solution of the problems which cannot be answered
by means of human pedigrees. In the case of the Cat the " trans-
mitting female " is visibly different from the non-transmitting, and the
most serious source of eiTor affecting the human data is thus avoided.

The character in the Cat which appears to correspond in its
inheritance with the sex-limited affections in Man is the orange colour
as contrasted with the black which corresponds with " normality." In
1904', in a short note on the subject, I concluded that in the Cat the
orange colour is dominant over black in the male, but only partially
dominant in the female, so that the female heterozygote is tortoiseshell.
The existence of sex-limited inheritance was at that time scarcely
known, but I mentioned the fact that among my collection of data
there was no case of an orange male mated with a black female giving
orange male kittens ; the females from such a mating were tortoiseshell,
the males black. Subsequent collection of further data has shown con-
clusively that the transmission of the orange colour by the male is
sex-limited, and the same result has been arrived at independently by
C. C. Little from his own experiments-.

In general, the results obtained with Cats are as follows : — an orange
male mated to a black female gives black male and tortoiseshell female
kittens ; in the converse cross, orange female by black male, the male
kittens are orange, the females tortoiseshell. The orange male thus
usually transmits orange to his daughters only, the orange female
transmits it to all her offspring of both sexes. A tortoiseshell female
by black male gives orange and black males, tortoiseshell and black

1 Proc. Camb. Phil. Soc. Vol. xiii. p. 35.

2 Scierice, May 17, 1912.

1() Sex-Limited Inheritance in Cats

t'ciiiales, showing that a female heterozygous for orange transmits the
orange factor to some kittens of both sexes. The colours cream and
blue appear to be dilute forms of orange and black, and to be inherited
similarly. In what follows I shall u.se the term "yellow" to include
orange and cream, "black" to include black and blue, tortoiseshell to
mean a mixture of yellow and black, and, when necessary, "bine-cream"
to describe the dilute tortoiseshell. The data given are derived almost
entirely from correspondence with numerous breeders, but include the
litters given by Little in the article in Science referred to. In the
information obtained from breeders I have made every effort to insure
that all inaccuracies are eliminated, but it is possible that a few
mistakes may have been included. Where I have good cause for
doubting the accuracy of a record, it has been omitted from the
tables.

A summary of the data collected with regard to the inheritance of
the yellow and black colours in Cats is given in the accompanying
table. It is noticeable that the divergences from equality of the two
sexes are similar in kind to those observed in the case of human sex-
limited diseases, where, however, as has been seen, it is doubtful
whether they are significant. In the offspring of yellow male mated to
black female there is some excess of females (61 % : 505/), as is also the
case among the offspring of men affected with one of the four diseases
mentioned previously; among the offspring of tortoiseshell females by
black males there is a considerable excess of males (07 (/" : 85 J ), as has
generally been found to be the case among the children of women who
transmit the diseases.

Another point of great importance is that there is evidence that
the sex-limitation of the transmission of the yellow factor by the male
is not absolute, for thirteen black or blue females are recorded from the
cross yellow male mated to black or blue female, five from yellow male
by tortoiseshell female, and three tortoiseshell females from one mating
of yellow male by yellow female. It should be mentioned that the
majority of the eighteen black or blue females from yellow sires were
blues, and it is possible that the simple explanation that sex-limitation
is not absolute, is not the. true one in every case. Apart from the
po.ssibility that the sex may have been wrongly recorded in .some
instances, it is not completely certain that a young kitten may not
sometimes be recorded as a blue when it is in reality a blue-cream.
The breeders of whom I have enquired on this point are not agreed ;
some say that blue-creams are always easily distinguished from blues at

L. DONCASTER

17

TABLES OF MATINGS.

I. Black or blue female x orange or cream male.
Offspring

Number

of matings

recorded

Orange

or

cream male

Black

or

blue male

Orange
or cream
female

Tortoiseshell

or blue-cream

female

Black

or

lilue female

25

—

46 + 3?

—

48

13

—

+

1 blue-cream i

—

— .

—

Note. Three blacks were of uncertain sex. In one mating of black ? x yellow ^ there
were produced two black ^ , three tabby j ; two tortoiseshell ? , and one tabby-
tortoiseshell ? (not included in the table).

II. Orange or cream female x black or blue male.

Offspring

Number Orange Black Orange Tortoiseshell Black

of matings or or or cream or blue-cream or

recorded cream male blue male female female blue female

6 20 — — 16 —

III. Tortoiseshell female x orange or cream male.

Offspring

Number

of matings

recorded

34

Orange

or

cream male

54 -fl?

-I- 1 tortoiseshell i

Black

Orange
or cream
female

blue male

38-1-3? 47-1-1

-t-2 smoke —

Tortoiseehell Black
or blue-cream or

female blue female

43 5

Note. Those marked ? were of uncertain sex. In addition, from three matings of yellow
i by " tabby- tortoiseshell " and " grey-and-orange " females, there were produced
four yellow s , three black or blue i , two tabby (j ; six yellow ? , two tortoise-
shell ? , one tabby ? .

IV. Tortoiseshell female x black or blue male.

Offspring

Number
of matings
recorded

22

+

Orange
or
cream male

35

1 tortoiseshell

•S

Black

or

blue male

29

f 2 smoke

Orange
or cream
female

Tortoiseshell

or blue-cream

female

21

Black

or

blue female

12

-1-2 smoke

Note. In addition, one mating of black (^ x "brown-orange-tabby" ? produced two
yellow c? , one black s ; one tabby ? ,

V. Orange or cream female x orange or cream male.

Offspring

Number
of matings
recorded

17

Orange Black Orange

or or or cream Tortoiseshell

cream male blue male female female

48 _- 40 3

blue female

Note. The three tortoiseshell females (one of them "blue with a cream patch") were
from the same mating (two litters),
Journ. of Gen. iii 2

IS Sex-Limited Inheritance in Cats

birth, others that mistakes may be made. In some cases at least these
blues grew up and seem to have been undoubted blues, and I know of
no case of two blues mated together giving creams, as should happen if
an apparent blue may ever be heterozygous for cream. In any case,
the few black females from yellow sires, about which there appears to
be no doubt, seem to prove that the sex-limitation of the transmission
of yellow by the male is not absolute.

Another point of considerable importance in connexion with the
comparison with the human cases is the ratio of the yellow to black
males in the offspring of tortoiseshell female x black male. As was said
above, in the offspring of transmitting females in the human cases, it is
generally believed that there is an excess of affected sons, but Lenz
regards this as due to the fact that a transmitting woman can only be
identified by having at least one affected son, and that this inevitably
raises the apparent ratio of affected to unaffected. When only large
families are considered, this excess disappears in the case of Colour-
blindness and Night-blindness, but remains in the Nystagmus and
Haemophilia totals, chiefly owing to the very great excess of affecteds
in certain pedigi-ees, but it seems doubtful whether even in these
affections it can be regarded as genuine. In the Cat, in which the
" transmitting female " (tortoiseshell) is visibly diffei-ent from the " non-
transmitting " (black), a small excess of " affected " (yellow) over black
is found (35 yellow : 29 black), but as in the human cases there is also
an excess of " transmitting " (tortoiseshell) over " non-transmitting "
(black) daughters, (21 : 12). The numbers are small, and further data
are required before they can be regarded as significant, but they give
no support to the suggestion discussed above that the excess of affected
males is due to partial coupling of the factor for the affection with a
sex-factor in the gametes of the transmitting fi'male.

One of the most interesting questions connected with the inheritance
of these colours in the Cat is the nature and origin of the rare tortoise-
shell males. Three of these are included in my records, one (a blue-
cream) from the mating yellow ^ with black % , one from tortoise-
shell female by black male, and one from tortoiseshell female by yellow
male. Two tortoiseshell males are also recorded from this last mating
in a note in Fur and Feather for May 10, 1912. I know of no other
case in which the male parent was known ; the few others that I have
met with have been produced by tortoiseshell females by unknown sires.
It has been shown above that there is reason for believing that the sex-
limitation of the transmission of yellow by the male Cat is not absolute,

L. DONCASTBR 19

since a small proportion of black females are produced from yellow sires,
and if this is so, there should theoretically be as many cases in which a
yellow male transmits yellow to his sons, as there are of his failing to
transmit it to his daughters. I know of no satisfactory record of a
yellow male mated to a black female having yellow sons, but suggest
that the tortoiseshell male is produced when, exceptionally, yellow is
transmitted by a yellow male to a son'.

Very little is known of how a tortoiseshell male transmits cplour-
factors to his offspring. The few breeders who possess them commonly
mate them to tortoiseshell females, in the belief that this is the most
likely mating by which to produce tortoiseshell male offspring, and I
know of only one mating with a black female, which is the one required
to test the matter thoroughly. In this one case the female was not
kept in confinement, so that although the pairing was seen, the
parentage of the kittens cannot be regarded as certain ; the only
recorded kittens were a black male and a tortoiseshell female. When
mated with tortoiseshell females, tortoiseshell males appear to behave
like yellows, giving yellow and black male, tortoiseshell and yellow
female offspring. Sir Claude Alexander, who has made many such
matings with one of his well-known specimens, writes of this tortoise-
shell male that with unrelated tortoiseshell females he " sires tortoise-
shells freely," but that with his own tortoiseshell daughters he gives
chiefly yellows, with only an occasional tortoiseshell female or a black.
Unfortunately, no record of the sex of these kittens appears to have
been kept ; no tortoiseshell males were produced.

The facts, then, as far as they are known, of the transmission of the
yellow colour in the Cat, may be summarised thus. In the female the
factor for yellow when homozygous produces orange (or cream when
dilute); when heterozygous produces tortoiseshell. In the male the
presence of the yellow factor normally produces orange (or cream), but
such males are not homozygous, for in general they transmit the yellow
factor to their daughters only. Exceptions to this rule, however, occur,
not apparently very rarely, for black females from yellow sires are
frequently recorded. Less common are tortoiseshell males, about which
little is known, except that they may apparently derive the yellow
factor from either parent, and that there is no recorded case of their
having tortoiseshell male offspring.

Until more data are collected, it seems of little value to attempt t<j
express these facts in a factorial scheme, for no means of testing such

' See also p, 21 below.

2—2

20 Sex-Limited Jnheritance in Cats

schemes are available. Certain points of importance, however, may be
considered. It has been suggested by Little^ that both the black and
yellow factors are sex-limited in the male Cat, and this suggestion is in
harmony with the fact that a female heterozygous for yellow is tortoise-
shell while the heterozygous male is yellow. For if the factor for black
were not sex-limited in the male, one would expect the male heterozygous
for yellow to be tortoiseshell as in the case of the female. If all males
containing the yellow factor were yellow, and if there were no exceptions
to the rule that the yellow male transmits yellow only to his daughters,
the case of the Cat would fall quite simply into the scheme which I
recently suggested for characters which appear to be dominant in one
sex and recessive in the other-. This suggestion was that one sex
(in the Cat and in Man the female) is normally homozygous for a
factor for " normality " (NN), for which the normal male is heterozygous
(Nn), and that N is constantly coupled with a sex-determining factor
for which the female is homozygous, the male heterozygous. In the
Cat the factor for normality would be the black determiner, which may
therefore be written B instead of N ; then if X is used for the sex-
factor, adopting the notation commonly used in America, a black male
would be BX.bx, a black female BX.BX. If then the yellow coloui- be
caused by the loss of the factor B, the yellow female would be bB.bX,
the tortoiseshell female BX.bX, and the yellow male bX.bx.

This scheme, however, has the obvious defect that it represents the
black male and the tortoiseshell female as of the same factorial composi-
tion as far as B and b are concerned (Bb in each case). An improvement
might be to represent the yellow flictor as a modification of B rather
than its absence; if the modified B which gives rise to yellow instead of
black be called Y, the yellow female would be YX.YX, the tortoise-
shell female BX.YX, and the yellow male YX.bx. Since Y is normally
coujjled with X, a yellow male would then normally transmit Y only
to his daughters, and since when he is mated with a black they would
receive B from their mother, they would be tortoiseshell, thus : —

yellow male YX . bx x BX . BX black female
gametes bx, YX BX, BX

black male bx . BX YX . BX tortoiseshell female.

' Scienc; May 17, 1912, 2 Joiinml of Genetics, i. 1911, p. 377.

L. DONCASTER 21

If now, exceptionally, Y became dissociated from X in the gametes
of the yellow male, and were thus transmitted to a son instead of a
daughter, the result would be

yellow male YX . bx x BX . BX black female
gametes Yx, bX BX, BX

tortoiseshell male Yx . BX b% . BX black female.

The female offspring would in this case be black, but heterozygous
for B instead of homozygous, and the male, since he contains both Y and
B, would be tortoiseshell.

I have one record of a tortoiseshell male produced by tortoiseshell
female by black male. The kitten died when only a month old, but
the breeder is confident that it was a male. If, as suggested, the
tortoiseshell male has the constitution YBXx, this could only arise in
this instance by the tortoiseshell mother ti-ansmitting YX, the black
father Bx ; the tortoiseshell male would then arise by the exceptional
association of B with x in the spermatozoa, instead of B with X'.

The scheme outlined above is satisfactory up to this point, but it
has two disadvantages. It involves the assumption that the occasional
black females produced from yellow sires are of different constitution
from normal blacks (BX.bX instead of BX.BX), for which there is no
direct evidence. The point could be tested by mating such a black
female with an orange male ; if the female wei-e heterozygous for B, it
would then give not only tortoiseshell females YX.BX, but also yellow
females of constitution YX . bX. This experiment has never, as far
as I know, been made. The second objection is more serious. The

1 There is an additional iiiece of evidence supporting the hypothesis that the factor B
is sex-limited. I have a record of tabbies being produced from yellow t? x black ? . Tabby
is known to be epistatic to black, but presumably can only appear in the presence of the
factor B. The yellow male which transmits the tabby factor does so to both sexes, the
male kittens which receive the tabby factor being brown tabbies, the females (only one
recorded) " tabby- tortoiseshells." It is clear therefore that the tabby factor is not sex-
limited. If, however, the black factor were present in the orange male, it is probable
that the cat would be tabby or tabby-and-yellow instead of yellow. If, therefore, a yellow
male can in some cases transmit the tabby factor, additional evidence is provided for the
assumption that the black factor is not present. Tabby tortoiseshell males, though very
rare, do exist, and are presumably tortoiseshell males which bear the tabby factor in
addition. One was exhibited at the Crystal Palace Show in 1912.

22 Sex-Limited Inheritance in Cats

tortoiseshell male should on this hypothesis be Yx.BX, and since B is
coupled with X, such a male should transmit B to his daughters and Y
to his sons. Mated to black he should have black female and tortoise-
shell male offspring ; mated to a tortoiseshell he should have tortoiseshell
and black daughters, yellow and tortoiseshell sons. Of the mating of a
tortoiseshell male with a black I have only one somewhat doubtful
record, but hope to be able to tost the matter with a tortoiseshell male
now in my possession ; in the several matings with tortoiseshell females
which are reported, no tortoiseshell males were produced — the tortoise-
shell male behaved as far as can be judged from the very incomplete
reports as if he were a yellow.

Otiier jjossible schemes might be suggested, for example that yellow
is due to an additional factor modifying B and usually coupled with it,
but occasionally becoming dissociated so that the yellow male would
transmit B to a female kitten and the modifying fiictor to a son. The
tortoiseshell male so produced, however, would still receive the modifying
factor from the male parent and B from the female, and it would be
expected, if these factors followed the ordinary rides of gametic coupling,
that they would show repulsion in the gametogonesis of the tortoise-
shell male, so that again the modifying factor would be transmitted to
the male offspring and tortoiseshell males would bo jjroduced as in the
previous scheme.

It seems clear, therefore, that the data at present available are
inadequate for the formulation of a factorial scheme of colour-inheritance
in Cats, which shall be in accord with previously known cases of sex-
limited inheritance and of gametic coupling in other forms. For any
one, however, who is able to undertake the work, the exjjeriments
required are fairly clear, and the importance of the case is increased by
the fact that the transmission of the yellow colour in the Cat is in close
agreement, both in its normal course and in the exceptions which occur,
with the transmission in Man of the sex-limited affections Colour-
blindness, Night-blindness, Nystagmus and Haemophilia. Although
so much has been written about the transmission of these diseases, no
agreement has been reached, and it seems probable that the inheritance
of colour in the Cat, with which definite experiments can be made, and
in which the " transmitting female " can be distinguished at sight from
the " non-transmitting," may provide the clue which the most comj)lete
human pedigrees would fail to yield.

In conclusion, I wish to tender my thanks to the following ladies
and gentlemen who have provided the information given in the

L. DONCASTER 23

preceding pages, and to many other correspondents who have helped
me in various ways to collect and verify my data.

R. Alder, Esq., Sir C. Alexander, Bart., W. 0. Backhouse, Esq.,
Miss W. Beal, The Hon. Mrs C. Behrens, Mrs Bennet, J. Bishop, Esq,,
J. L. Bonhote, Esq., Miss Cope, Miss A. M. Drury, The Misses Elli<ttt,
Mrs Forsyth Forrest, Miss F. E. Frowd, Mrs M. M. Hall, Mrs Kennaway,
Miss H. Lea, Mrs L. Maxwell, Mrs Norris, Mrs F. Owen, Mrs Simner,
Mrs Slingsby, Mrs Mackenzie Stewart, Miss H. M. Tawse, Miss Turnhill,
Miss Veevers.

Cambridge,
January 28^ 1913.

FIVE GENERATIONS OF CONGENITAL STATIONARY
NIGHT-BLINDNESS IN AN AMERICAN FAMILY.

By H. H. NEWMAN.

{Fi'om the Hull Zoological Laboratory, University of Chicago.)

In his book Mendel's Principles of Heredity, BatesoD, discussing
the laws of sex-limited inlieritaiicc in man, says that " the only one of
the notoriously sex-limited conditions which is available for testing the
rules is colour-blintiness. Haemo2:)hilia and Gowers' disease are too
fatal and night-blindness is too rare and too little known." It is the
purpose of the present paper to add a chapter to the present meagre
literature on the inheritance oF night-blindness'. The data herewith
presented were obtained in collaboration with Miss Elizabeth L. Brown,
who has had exceptional opportunities for getting at the facts since she
is a member of the affected family and has seen and tested the majority
of her living relatives. Those whom she has been unable to observe
have freely communicated the facts to her. I have been in correspond-
ence with Miss Brown for considerably over a year and have directed
her investigations. Every effort has been made to eliminate all sources
of error and to obtain all information that might be of value. There is.
still some doubt as to the exact character of the defects associated with
night-blindness, but we can vouch for the accuracy of the pedigree as
given in the diagram in so far as the major defect, night-blindness, is
concerned. Errors have been eliminated from the pedigree largely
through the use of a questionnaire suggested by Mr Nettleship, who
has been kind enough to express a personal interest in the progress of
the investigation.

' A very recent paper by Nettleship has just come to my attention in which a case of
night-bUndness similar to that here dealt with is discussed. The paper is : "A Pedigree
of Congenital Night-Bliiidness with Myopia." Ophth. Soc. Trans. Vol. xxxii. 1912.

26 Congenital Nig/if -Blindness

The association of families dealt with in this paper is confined to
the State of Texas, "but o*?-iginated in the State of North Carolina.
Whether or not the strain can be traced back to an English origin I am
unable to ascertain. It would be interesting to find that the present
line is a branch of one of those investigated by Mr Nettleship. In order
that it may be made possible to trace such connection, if it exist, I shall
give the full names, where obtainable, of all individuals concerned.

Especial interest attaches to the present pedigree first because it
appears to follow a formula hitherto nndescribed for night-blindness
and second because the major defect is found so generally associated
with much less rare disease of the cornea known as Pterygium, which
is not sex-limited but more pronounced in males than in females.
Strabismic and myopic conditions appear to be in some way tied up
with night-blindness and are also sex-limited.

The peculiarities in the descent of the various sex-limited characters
of man are given by Bateson as follows :

1. They affect males much more commonly than females.

2. They may in certain cases be transmitted by affected males, but
are not transmitted by unaffected males.

3. They are nevertheless transmitted by the unaffected females.
Apparently normal women, sisters of the affected males, thus may
transmit the condition to some of their sons.

The case herewith described falls in line with none of these
schemes of inheritance but would demand a fourth formula somewhat
as follows :

4. They are not transmitted directly by males, either affected or
unaffected, hid only by the imaffected daughters of uff'ected males to some
of their sons.

That this method of inheritance was recognized by Bateson but
inadvertently omitted is shown in a note facing page 230 of liis book, in
which he corrects an error regarding the inheritance of colour-blindness.
He says ;

"In haemophilia and night-blindness there are cases of the direct
descent of the condition from father to son in families showing ordinary
sex-limited descent, but as regards colour-blindness, though there are
instances of the direct descent from father to son, it is now obvious that
in all of them the affection was introduced by the normal-sighted
mother also. Apart from these there is no case of a colour-blind man
having a colour-blind son known to mo. On the contrary, from the
records, fir the most part communicated by Mr Nettleship, there is

H. H. Newman 27

now good evidence as to the condition of 23 sons of colour-blind men
and all are normal (besides 15 more said to be normal on less ceitain
authority). Such a fact is very significant."

That colour-blindness is not the sole optic affection following this
interesting inheritance formula is shown in the present case where
night-blindness is seen to follow exactly the same course.

Following the method adopted by Nettleship and others I shall first
give a pedigree showing the inheritance of night-blindness, dealing
with the less certain accompanying affections in connection with the
descriptions of individual cases. The five generations will be taken up
serially, beginning with the oldest, and each individual will be referred
to a number shown in the diagram.

I, 1. Joel Bryan of North Carolina, a Quaker, probably of English
extraction. No definite information as to whether or not he was night-
blind.

I, 2. Wife of Joel Bryan. No information obtainable concerning
her maiden name or antecedents.

I, 3. Thomas W. Vause of North Carolina, a physician, a first
cousin (?) of Joel Bryan, but considerably younger than the latter. As
a young man Mr C. D. Uzzell carried on a considerable correspondence
with Dr Vause with reference to the night-blindness with which both
were affected. There is no doubt then that niglit-blindness was of
earlier origin than this generation.

II, 1. Margaret Bryan Uzzell, daughter of Joel Bryan, is the only
issue of the latter that it has been possible to bring to light. Several
years after her marriage to Mr Uzzell in North Carolina she moved to
Texas and was the ancestor of the present Texas night-blind connection.
She had normal vision but transmitted night-blindness to three of her
four sons. There were in addition five normal daughters.

II, 2. Mr Uzzell, husband of Margaret Bryan Uzzell, had normal
vision, as is testified by his son, Mr C. D. Uzzell.

III, 1 — -11. Issue of II, 1 and 2. No other pregnancies known to
Mr C. D. Uzzell.

1. J. C. Uzzell, dead, night-blind to the same extent as C. D.
Uzzell (5). He seems to have left no progeny.

2. Mrs Haynie, dead. There is no record of night-blindness in her
or in her progeny.

3. Mrs Bayers, dead. Normal and normal progeny.

4. Mrs C. U. Carson, aet. (1912) about 77, normal vision and
normal progeny.

28

Congenital Niglit-BUndness

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H. H. Newman 29

5. C. D. Uszell, aet. (1912) 74, night-blind from infancy, "short-
sighted" (probably myopic) and has suffered considerably fi-om Pterygium,
having been rather recently operated on for the latter with apparently
beneficial results. He is the father, by his normal and unrelated wife,
of six normal sons and four normal daughters and the grandfather
of nine normal and ten night-blind grandsons and of eleven normal
grand-daughters. Mr Uzzell has taken a lively iuterest in the present
investigation and has furnished some of the most important information.
A quotation from one of his letters should prove interesting: "Night-
blindness and near-sightedness were transmitted to our family on the
maternal side, but personally I never knew of but one case in a
preceding generation, and he was a (second) cousin of my mother. He
was a good physician and somewhat of a zoologist. I used to have an
extensive correspondence with him, hojDing thereby to find some way to
remedy ray own defective vision, but he was unable to assist me in any
way. My mother, who was perfectly normal in vision, had four male
children, three of whom were affected just as I am, and one that was
not. (He failed in this letter to mention that he had five normal
sisters, but this fact was brought out later.) All my children have
perfect eyes. This trouble does not seem to be transmitted regularly,
but it seems to fall on alternate generations, so my mother told me. It
is also transmitted from the father through the daughter to the grand-
sons. I have never heard of a girl in the family connection who did
not have perfect sight. The children of both my sons (referring only to
those with families) have perfect sight, but the eyes of my daughters'
sons are nearly all affected, one daughter having seven sons, five of
whom are affected. My other three daughters have each one son and
each is affected. (Reference is made to living sons only. One grand-
son V, 30 has been born since the receipt of this communication.)"
Miss Elizabeth L. Brown (V, S) writes of C. D. U., her grandfather :
" Grandfather (III, 5) was unable to see the moon and stars until
he was about thirty. He sa3's the defect grows less and less as he
grows older. He has had much trouble with a growth similar to
cataract, which grows in the inner corner of the eye and gradually
spreads toward the centre of the eye with advancing years. (This con-
dition has been diagnosed as Pterygium, a progressive formation on the
cornea, beginning as a pinguicula located at the nasal angle of the
cornea and overgrowing the cornea as a fold or wing of tissue, usually
highly vascular in character.) It sometimes becomes inflamed. Opera-
tions have relieved the condition to some extent."

30 Congenital Night-BUndness

Note: — The parentheses in both of the above (juotations are my
own.

6. Normal wife of C. D. Uzzcll.

7. Normal wife of E. B. Uzzell.

8. E. B. Uzzell, aet. about 70, brother of C. D. U., is night-blind,
myopic and has suffered greatly with Pterygium. He was almost blind
a few months ago, according to his own statement, but has been partially
relieved by an operation. He writes that all of his children and those
of his sons are normal so far as night-blindness is concerned, but that
the sons of his daughters show a considerable percent, of night-blindness
and other defects of the eyes. He has five normal sous and two normal
daughters, both of whom have night-blind sons.

9. Mrs N. Carson, dead, normal in visiou. There is no known case
of night-blindness in her progeny.

10. Mrs L. Doyle, living, normal, with normal progeny.

11. William Uzzell, living, the only normal male of the childship.
None of his progeny or descendants exhibit night-blindness.

IV, 1 — 16. Issue (minus the marriages) of III, 5 and 6. No other
pregnancies.

1. First-born male child. Evidently died in infancy as no name
was given.

2. H. W. Uzzell, dead, normal. Left no progeny.

3. D. K. Uzzell, aet. 43, normal so far as night-blindness is con-
cerned, but his eyes "are in very bad condition from cataract" (Pterygium
is the diagnosis here). Three of his four living sons show symptoms of
Pterygium more or less advanced. His two daughters, now dead, were
normal.

4. Normal wife of D. K. Uzzell.

5. Mrs U. S. Brovm, aet. 41, normal so far as night-blindness is
concerned, but has the most pronounced case of Pterygium among the
female members of the connection. She has had two male and five
female children. Both sons, one of whom is now dead, night-blind.
The daughters, two of whom are living, all normal. Miss E. L. Brown,
who has collaborated with the writer in this investigation, is V, 8, the
eldest child of Mrs (J. S. Brown.

6. Normal husband of Mrs U. S. Brown.

7. Mrs Hohdy, aet. 39, normal. She has seven sons living, five of
whom are unquestionably night-blind and two are normal. No
Pterygium apparent.

8. Normal husband of Mrs Hobdy.

H. H. Newman 31

9. Normal wife of M. W. Uzzell.

10. M. W. Uzzell, aet. 37, normal so far as night-blindness is con-
cerned, but has an incipient case of Pteiygiiiin, the least troublesome
case among the males of the third and fourth geueratious. He has only
one child, a normal son, aged 3.

11. Normal husband of Mrs E. F. Floyd.

12. Mrs E. F. Floyd, aet. 35, normal so far as oight-bliuduess is
concerned, but shows a moderate development of Pterygium. She has
had two uight-blind sons, one of whom is now dead, and two normal
daughters. The living son and one of the daughters are twins.

13. C. D. Uzzell, Jr., aet. 32, normal so far as night-blindness is
concerned, but with a pronounced case of Pterygium. He is unmarried.

14. J. U. Uzzell, aet. 30, normal so far as night-blindness is con-
cerned, but with Pter3gium obvious. He is unmarried.

1-5. Mrs Hurst, aet. 27, normal. She has two sons, one certainly
night-blind and the other too young to exhibit the defect. She has also
two normal daughters.

16. Normal husband of Mrs Hurst.

IV, 17 — 25. Issue (including normal mates) of III, 7 and 8. No
other pregnancies.

17, 18, 21, 22. Normal sons of E. B. Uzzell. They are all married
and have none but normal issue.

19. Mi's Montgomery. Normal. Has two uight-blind sons and
one normal son, and one normal daughter.

20. Normal husband of Mrs Montgomery.

23. Mis Morton. Normal. Has one night-blind son.

24. Normal husband of Mrs Norton.

25. Normal unmarried son of E. B. Uzzell.

V, 1—7. Issue of IV, 3 and 4.

1. Irvin Uzzell, aet. 19, normal so far as night-blindness is con-
cerned and without Pterygium.

2. Wason Uzzell, aet. 17, normal so far as night-blindness is con-
cerned. Has had some trouble with Pterygium.

3 and 4. Jeiuel and Minter Uzzell, dead. Both were normal.
5 and 6. Lester, aet. 11, and Bouglas Uzzell, aet. 9, normal so far as
night-blindness is concerned, but both have incipient Pter^'gium.

7. Female infant, died in infancy.
V, 8—14. Issue of IV, 5 and 6.

8. Elizaheth. L. Brown, aet. 21, normal. Miss Brown is a member
of the present senior class at the University of Texas and has made a

32 Congenital Nkj Id -Blindness

speciality of Zoology. Without lier aid it would have been practically
impossible for me to have gathered the data for this paper.

9. Henri/ W. Brown, dead, was unmistakably night-blind and
showed the usual associated defects.

10. Aline Brown, dead, normal.

11. Leona Brown, aet. 14, normal.

12. Marjory Irene Brown, dead, normal.

13. Stella K. Brown, aet. 9, normal.

14. Walter S. Brown, aet. 4, night-blind, and slightly strabismic.
Of him Miss E. L. Brown, his sister, writes: "Brother cannot see
anything out of the light at night, except that on bright nights he can
see anything between him and the sky. I took him out of doors one
night and noted that he could see the wind-mill, trees, houses, fences,
etc., but at the same time he could not distinguish his express wagon,
his kitten or other objects on the ground. He tries to count the stars
and the lights in other windows, so I am sure that he is less night-blind
than some of the Floyd and Hobily children. He is not short-sighted
at all, but his eyes do not focus together. This was more noticeable
when he was a bab}', but he has outgrown it until it is only noticeable
when he looks up."

V, 1.5—21. Issue of IV, 7 and 8.
1-5. Drew Hohdy, aet. 19, normal.

16. Joltn Hohdy, aet. 10, night-blind, myopic and strabismic.

17. Walter B. Hobdy, aet. 13, night-bliod, but without any other
optic defect.

18. Maurice Hohdy, aet. 11, normal.

19. Monroe Hohdy, aet. 8, night-blind, myopic and strabismic.

20. Alton Hohdy, act. 6, nigiit-blind, myopic and strabismic. Was
tested as follows : Turns head to one side when reading, holds book less
than 8 inches from the eye.s.

21. Vernon Hohdy, aet. 4, night-blind, but without the other
associated defects.

V, 22. Minter Uzzell, Jr., only issue of IV, 9 and 10, normal,
aet. 3.

V, 23—26. Issue of IV, 11 and 12.

23. Enid Floyd, aet. 10, normal.

24 and 2.5. Ira and Irma Floyd, twins, aet. 5. Irma is normal.
Ira is night-blind, myopic and strabismic. Miss E. L. Brown writes of
him as follows: "He holds objects as small as a pin about five inches
from his eyes. Being too young to read I could not tell how far from

H. H. Newman 33

his eyes he would need to hold a book. The right eye looks straight
ahead, but the left does not focus with the right. In looking closely at
objects he turns his head to one side and looks with the left eye. He
can hold his hand over the left eye and see well with the right. He is
being treated for this trouble by an oculist who claims that he will be
able to see at night when he has finished with him."

26. Edward T. Floyd, dead. Lived long enough to exhibit sure
signs of night-blindness. No facts about associated defects.

V, 27—30. Issue of IV, 1.5 and 16.

27. Floyd Hurst, dead, normal.

28. Mildred Hurst, aet. 6, normal.

29. Jack Hurst, aet. 4, night-blind, but without other defects.

30. Marvin Hurst, aet. 8 months (will be nearly 2 years when this
paper is published. All the ages used in paper apply to the Autumn
of 1911). He now shows unmistakable signs of night-blindness, but
no other defects.

V, 31—34. Issue of IV, 19 and 20.

31. J. L. Montgohiery, aet. 24, normal.

32. R. D. Montgomery, aet. 21, night-blind and myopic (wears
concave glasses).

33. H. L. Montgomery, aet. 18, night-bliml, myopic. Mr C D.
Uzzell (III, 5), his great-uncle, says of him : " His eyes are in terrible
condition. He can hardly see in the daytime. If he starts to go
through a door without his glasses he is as apt to miss it as to go
through it."

34. Evolyn Montgomery, aet. 8, normal.

V, 35. Issue of IV, 23 and 24. Name and age unknown. Night-
blind.

Discussion.

Tlie probable mechanism of the inheritance of night-blindness.

It has been shown by Guyer ('10) that there is a bivalent A'
chromosome in the spermatogonia of man and that two classes of
spermatozoa are formed in equal numbers, half with an X element and
half without^ In the light of the investigations of E. B. Wilson and
others it seems practically certain that the sex of man is in some way

' Some of the details of Guyer's results have been disputed, but need not be discussed
here.

Journ. of Gen. iii 3

34 ComjenUal Niyht-BUndness

dependent upon or associated with the presence or absence of the
X chromosome in the spermatozoa. Regarding the ova as homozygous
and the spermatozoa as heterozygous for sex, we may explain the
inheritance formula of night-blindness by assuming that the character
is a Mendelian recessive, which has probably arisen by the loss of some
factor necessary for normal vision, and that the defect has its seat ni
the X element of the affected male. Probably night-blindness arose in
a male as a mutation.

On this assumption eggs fertilized by spermatozoa with the A'
element will develop exclusively females, and therefore the X element
of the affected father, together with whatever defects or peculiarities
may be associated with it, is transmitted only to the female offspring.
This explains why all of the sons of affected men are normal. The
daughters, however, all inherit the factor for night-blindness, but fail
to show it because it is recessive to the normal character carried by the
normal X element of the normal mother. The maturation of the ova
of these night-blind carrying daughters will result in the formation of
gametes of two kinds in equal numbers, half with the normal A' element
and half with the defective A element (= x) carrying night-blindness. If
these daughters are mated with normal men we should expect equal
numbers of male and female offspring, half of each of which would
contain the factor for night-blindness. The females of course would
always appear to be normal, but half of the sons would be night-blind.
The following diagram (Fig. 2) will illustrate graphically the inheritance
mechanism just outlined.

This diagram is a modification of that used by E. B. Wilson ('11)
in explaining his idea of the probable mechanism of the heredity of
white eyes of Drosophila. He states that the same type of explanation
may be applied to the inheritance of colour-blindness in man. Since
this case of night-blindness is inherited precisely as is colour-blindness,
there can be little hesitation in adopting the scheme outlined above
for its explanation.

Wilson points out, furthermore, that " any recessive mutation should
exhibit sex-limited heredity when crossed with the normal or dominant
form, if it be due to a factor contained or omitted from the X element."
It is now known that a number of human traits of an abnormal or
defective nature are inherited in the fashion exemplified by the present
pedigree. Davenport ('11) gives the following list of such characters:
multiple sclerosis, hereditary atrophy of the nptic nerve, colour-blind-
ness, myopia, ichthyosis, muscular atrophy, haemophilia. Probably

H. H. Newman

35

this mode of inheritance will be found, in the near future, to be unich
more common than now appears to be the case.

It is an interesting thought that a specific defect in the adult may
be due to an equally specific defect in a single chromosome of the germ
cell. Evidently any defect in the X element, assuming that the latter
is the seat of the factors responsible for the various defective ^conditions
listed above, must be as distinct and specific as is the condition for
which it becomes responsible in the adult. One might, on the other

Zygotes

Gametes

Zygotes

Gametes X

Zygotes X X x X X

Fig. 2. (Explanation on p. 38.)

hand, conceive germinal deficiency as due to a generalized metabolic
weakness of the X element, and might assume that the particular form
of adult deficiency arising from this situation depends on the specific
peculiarity of the particular line of germ plasm in which the germmal
deficiency arose. In this way one simple type of germinal defect
might in one family result in a complex of optic defects and in another

as haemophilia or musculs.? atrophy.

3—2

36 Conf/eiiital Ni(//if- Blindness

Ah-lioLigh the explanation here ofifered to show how night-blindness
is probably inherited in this instance is on the whole rather satisfying,
it does not completely accord with the data shown in the pedigree.
Perhaps the most serious tliscrepancy is encountered in connection with
the apparent non-inheritance of the capacity to transmit night-blind-
ness on the part of the sisters of night-blind men. According to
expectation there should be in F,, or F-^, assuming that there has been
no consanguineous marriage, ecjual numbers of night-blind carrying and
of normal females, but the pedigree seems to show that none of the
progeny of the five married sisters of C. D. Uzzell and E. B. Uzzell
(night-blind males of the third generation) show a trace of night-
blindness. Whether sisters of night-blind males can transmit the defect
will doubtless be determined in the present generation. If it should
prove that night-blindness cannot be inherited for two consecutive
generations in the female line, it would be a fact of unusual significance.
It would seem to be a case in which a prolonged association of the
defective X element with the normal X element in the germ plasm of
two generations of females might serve to correct tlie defect in the
former and thus eliminate night-blindness from these particular lines.
The value of this idea depends on whether or not the findings in the
pedigree are substantiated by future events.

In common with other writers who have reported pedigrees of this
sort I find a serious discrepancy in the ratio of males and of females,
and in that of afrecte(is and unatfecteds. Of the 36 offspring of the
night-blind carrying daughters of tiight-blind men, 22 ai-e males and
14 females instead of the expected IS : 18. Perhaps this discrepancy
is within the range of probable error. But the fact that of these
22 males 17 are affected and only o unaffected can hardly be so
interpreted. An excess of afi'ecteds, however, is found quite generally
in haemophilia and in certain less known inherited affections, and it
may well be, as Bates(}n suggests, that in these cases there is present
some unknown disturbing factor that distorts the normal Mendelian
ratios.

The association of tdght-hlindness luith other optic defects.

Nettleship and othei-s have noted that the sex-limited type of
night-blindness is almost invariably associated with myopia. He even
finds it sometimes associated in inheritance with certain mental defects.
The present pedigree is in agreement with the conditions described by

H. H. Newman 37

Nettleship with respect to rayopia, and I find strabismus' almost as
invariable an accouipaniment of night-blindness. An expert ophthalmo-
logist would doubtless detect several other well-defined affections, which
almost certainly are present in those cases where a near approach to
complete blindness is described, for, as I understand it, complete
blindness is never the result of eitiier night-blindness or any of the
associated conditions known to exist in this family.

The frequent occurrence in tiie pedigree of the minor defect
Pterygium^ seems to be somewhat unusual. The defect is evidently
strongly inherited, probably as a Meudeiian dominant, since it is neither
sex-limited nor necessarily associated with the recessive night-blind
complex. Peculiarly enough, however, although not strictly sex-limited,
it occurs more commonly and assumes a moie serious aspect in males
than in females.

It would appear then that night-blindness, myopia, strabismus (and
possibly some other conditions that have not been diagnosed) form a
complex due to a single sex-linked factor and that whether one, two
or all of the conditions appear simultaneously or successively in an
individual may depend upon cooperating or counteracting factors in
the germ plasm of the normal parent, producing an intensification or a
dilution of the somatic expression of the germinal defect.

In conclusion it may be well to point out that, if we follow Wilson's
suggestion, it is no longer admissible to interpret such conditions as
colour-blindness and this type of night-blindness as due to a factor
dominant in males and recessive in females, for it seems pretty well
established that such characters are really recessive and only express
themselves, after the manner of recessives, in the absence of the
dominant allelomorph.

' Nettleship suggests that strabismus is due to the necessity the affected are under
of bringing the book, etc. very close to the eyes, and is unrelated with night-blindness as
such.

^ Nettleship is o£ the opinion that Pterygium as such is not inherited but that there is
merely a congenital peculiarity of the mucous membrane predisposing to Pterygium. The
defect itself is commonest in people whose eyes are constantly exposed to glare, dust and
irritants over a number of years. The glare and dust of Texas are characteristic elements
of the climate.

38 Congenital Nir/Jif-BUndness

LEGENDS OF FIGURES.

Fig. 1. Pedigree showing the inheritance of night-blindness in a Texas family. The
black symbols are the affected members. The Eoman numerals on the left refer to
generations.

Fig. 2. Diagram to show the possible mechanism of the heredity of night-blindness.
The normal accessory chromosome is represented by X ; the abnormal accessory
chromosome by .t, which is assumed to carry the factor for night-blindness. In the
first generation x is in the male line ; in the second generation it always crosses over
to the female line and is dominated by A' ; in the third generation it should be equally
distributed between the male and female lines. A minus sign ( - ) indicates the
absence of the X or .r element.

LITERATURE.

1. Bateson, W. Menders Principles of Heredity. Cambridge, 1909.

2. Davenport, C. B. Heredity in Relation to Eugenics. New York, 1911.

3. GuvER, M. F. " Acces.sory Chromosonie.s in Man." Biol. Bull. Vol. xix. 1910.

4. Nettleship, E. "A History of Congenital Stationary Night-blindness in

Nine Consecutive Generations." 0}}hth. Sac. Trans, xxvii. 1907.

5. — _. "Three new Pedigrees of Eye Disease." lldd. .xxvrii. 1908.

ON STERILE AND HYBRID PHEASANTS.

By GEOFFREY SMITH, M.A.,

Fellow of New College, Oxford;

AND Mbs HAIG THOMAS, F.Z.S.

The material for this study consists of four abnormal or " mule "
pheasants, which on dissection proved to be females with aborted
ovaries, and a series of hybrids between Reeve's pheasant and the
Japanese pheasant and Formosan variety of the Chinese pheasant bred
by Mrs Haig Thomas and handed over to me for histological examina-
tion. Mrs Haig Thomas has also put at my disposal some valuable
statistics derived from her extensive experiments in breeding hybrid
pheasants of various kinds, which throw an interesting light on the
disturbance of the sex-ratio through hybridisation. The examination
of the plumage of these birds was made by Mrs Haig Thomas, and
I have also had access to her records relating to their behaviour, change
of plumage, etc., which occurred while they were living in her aviaries.
This paper is, therefore, a joint production, in which Mrs Haig Thomas is
responsible for the breeding experiments and myself for the theoretical
interpretation. The facts brought out by the study of these various
sources throw light on three distinct problems which may be treated
separately, first, the cause of the sterility of the hybrids in so far as it
is elucidated by a histological examination of the reproductive tissues,
second, the assumption of partial male plumage by the sterile females
and the correlation of this abnormal plumage with atrophy of the
ovary, thirdly, the preponderance of males over females in hybrid
pheasants and the question as to how far this preponderance is due
to a selective death-rate, and at what period the incidence of this
selective death-i-ate occurs.

40 Sterile and Hijlmd Pheasants

1. The cause of the sterility of the lii/brids.

In dealing with some sterile male hybrid pigeons in Part 9 of "Studies
in the Experimental Analysis of Sex," Q. J. M. S. Vol. i.viii. 1912, it
was shown that the point at which the formation of the spermatozoa
became abn(jrmal was at the synapsis preceding the first maturation
division. It was proved that the chromosomes in this division failed to
form the normal synaptic pairs but were distributed unequally on the
spindle, and it was also found that in the majority of cases the secondary
spermatocytes failed to divide again but went straight on to form sper-
matozoa of double the normal size, while in some cases deformed
spermatozoa with chromatic beads were produced.

Two sterile male pheasant hybrids produced by crossing Reeve's $
with versicolor j/ and Reeve's ? with Formosan ^^ have been inves-
tigated. The birds were killed in June, when the former was rather
moi-e than three years old and the latter two, and dissection showed
that the reproductive organs were anatomically normal. The vesiculae
seminales of the Reeve's by versicolor male were full of spermatozoa,
which moved actively when placed on a slide, though the presence of a
very large proportion of deformed spermatozoa was at once apparent.
In the other male the vesiculae were empty except for a certain amount
of cellular debris which appeared to have resulted from the breaking-
down of deformed spermatozoa. Smear preparations of the spermatozoa
from the first male, .stained with iron-haematoxylin and eosin, brought
out a condition of affairs very similar to that which was described for
the hybrid pigeons in Part 9, except that in this ease there was a much
greater preponderance of deformed spermatozoa. On comparing the
smear with similar preparations made from some normal male Phasi-
anus torquatus, it was apparent that the same irregularity in size of
the hybrid's spermatozoa occurs as in the case of the hybrid pigeons.
Figs. 1 and 2 on Plate I depict the normal (Fig. 1) and the hybrid
spermatozoa (Fig. 2), and it will be seen that many of the latter are
about double the normal size, while others are smaller than the normal.
In addition to these abnormal!}' sized spermatozoa, a few of the deformed
spermatozoa are drawn, but as a matter of fact in this hybrid these
deformed spermatozoa are greatly in excess of the rest.

The examination of sections of the testes of the hybrids brought
out that, just as in the hybrid pigeons, the spermatogonia are perfectl}^
normal in appearance, but all the subsequent stages show degenerative
changes in the nuclei of a more pi-onounced kind than was found in

Geoffrey Smith and Mrs Haig Thomas 41

the hybrid pigeons (Fig. 3, Plate I). The typical appearance of the
testicular tubes is shown in Fig. 3, Plate I, and it will be seen that
after the spireme stage of the primary spermatocytes, the cells appear
to have fragmented irregularly, and the chromatin in them appears as
intensely staining irregular masses which assume branching and beaded
threads. These degenerate fragments represent the abnormally shaped
spermatozoa which make up the greater number of the spermatozoa
found in the vesiculae seminales, and the method of their formation
appears to have consisted in a fragmentation of the spermatocytes after
the abortive initiation of the reducing division.

The occurrence in the vesiculae of a certain number of normally

shaped spermatozoa, though mostly of abnormal size, would seem to

indicate that a certain number of the primary spermatocytes did divide

in a less degenerate manner earlier in the year, but I cannot find any

-evidence of this occurring in the sections.

It will be seen from this account that degeneration sets in just at
the time when the synaptic pairing of the chromosomes should follow
the formation of the spireme in the primary spermatocytes, and this
agrees with the account given for the hybrid pigeons, except that the
degenerative process sets in rather earlier in the case of the hybrid
pheasants. This earlier degeneration in the latter case accounts for
the far greater proportion of deformed spermatozoa found in the
pheasant compared to the pigeon. Besides the two sterile males, three
sterile females from the same crosses, viz. two produced by Reeve's
$ X Formosan </ and one by Reeve's ? x versicolor- </", were dissected.
The Reeve's x Formosan hybrids were rather over two years old, and
had never laid eggs ; the Reeve's x versicolor female was three years
old and had likewise never laid.

The 'Reeve s X versicolor female (M) did not show any marked
assumption of male characters in the plumage, except in the pattern
of the interscapulars which showed a trace of the male Reeve's type.
There was a very small ovary which however contained some fair-sized
oocytes, and the oviduct was small and undeveloped. Unfortunately
the ovary of this bird was lost, so that no report can be made on its
histological structure. Of the two Reeve's x Formosan females, one,
which will be referred to as B. showed very slight assumption of
Reeve's male pattern in the interscapulars. The ovary consisted of
two nodules of tissue about 5 mm. in length, highly vascularised
and of a dark red colour. Sections of these nodules showed no trace of
oocytes, the whole of the tissue consisting of nests of interstitial cells,

42 Sterile and Hybrid Pheasants

with a good deal of stroma and fibrous tissue (Figs. 4 and 5, Plate I).
In places small cysts could be seen surrounded with a fibrous capsule
and containing a fatty fluid (Fig. 5, cy., Plate I). In every respect the
sections through this degenerate ovary resemble the Figures 19, 33, 34
and 39 given by Prof Poll for the ovaries of duck hybrids (1).

This bird was otherwise peculiar in that the body was laden with
subcutaneous and visceral fat to an extraordinary extent, and also in
possessing two very large hypertrophied oviducts.

The second Reeve's x Formosan female ((?) showed a very striking
assumption of male plumage, both in pattei'n and colour of the neck,
breast, interscapular and tail feathers. The ovary was still more
degenerate in structure than in the foregoing, consisting of some
minute nodules about 1 mm. across, which showed the same histological
structure as the other female. The oviduct in this female- was very
small and undeveloped. It may be noted that all the sterile females
were of an unusually large size, a character which may be regarded as
a male one.

The histological structure of the degenerate ovaries of these
females agrees in every way with the excellent descriptions which
Prof Poll has given for hybrid ducks; we see the same inability of the
oocytes to grow normally, and the consequent composition of the ovary
by interstitial, stroma and fibrous cells.

The interesting enquiry, which is not answered by Prof Poll, is to
determine whether the degeneration of the ova can be ascribed to the
same cause as that of the spermatozoa, viz. to the inability of the
chromosomes to go through the synaptic phases. At first sight such
an interpretation seems absurd, since degeneration of the ova occurs in
these hybrids long before jjolar-body formation when the maturation
divisions take place. But as I pointed out in a footnote in a previous
paper (Q. J. M. S. Vol. LViii. p. 166), there is evidence that in the
female sex a precocious temporary synapsis occurs in the young germ-
cells before growth of the oocytes begins, and that this may take place
in the quite young organism or even in the embryo. Since the publica-
tion of the above-mentioned paper, Mr Jenkinson has collected a great
deal of information on this point, and it seems clear that in the female
sex throughout the animal kingdom this temporaiy synapsis of the
chromosomes in the young oocytes before growth begins is of universal
occurrence. Furthermore Mr A. D. 8prunt has worked through the
early stages of developement of the ovary in the chick (Galliis), and he
finds that the typical synaptic phases occur in the embryonic ovary

GrEOFFREY Smith and Mrs Haig Thomas 43

between the seventeenth and nineteenth day of incubation. After
synapsis the chromosomes separate and the nuclei pass back into the
vegetative phase until polar-body formation. Now it is perfectly clear
from the foregoing facts that since in the normal female it is necessarj-
for the oocytes to undergo the synaptic phases before they can store
yolk and grow up into mature oocytes, we must conclude that in the
case of the sterile hybrids this early synapsis was initiated, but owing
to the incompatibility of the chromosomes it failed to take place
normally, with the result that the oocytes failed to grow.

The fact, therefore, of this early synapsis in the female germ-cells,
gives us a complete explanation of why the germ-cells degenerate in
the female at an apparently earlier stage than the germ-cells in the
male. In reality the germ-cells begin to degenerate and become ab-
normal at exactly the same phase in both male and female hybrids, viz.
at the synaptic phase, so that the generalization that the sterility of
hybrids is due to the incapacity of the chromosomes derived from two species
to form the normal synaptic pairs applies to both sexes of hybrids, and is
probably of universal application.

2. The assumption of male characters by sterile females.

The sterile females examined under this head include four Common
Pheasants (A, G, B and E), which were probably all crosses between
P. torquatus and colchicus ; a sterile pure-bred Formosan variety of
P. torquatus (K) ; a female hybrid between Reeve's $ and versicolor
J" (M), and two hybrids between Reeve's $ and Formosan J' (B
and G).

The Common Pheasants A, G, D and E will be dealt with first.
These were all young birds, shot probably in their first winter, and
dissection revealed an identical condition in all of them, viz. entire
absence of any visible ovary or gonad of any kind, and the presence of
a very small, narrow oviduct on the left side which had its opening into
the cloaca occluded, and the other end narrowing down to a fine fila-
ment. I was unable to detect any trace of germinal tissue in any of
these birds; otherwise they appeared healthy and normal. All four
birds showed male characters in the plumage to slightly varying
extents. In all of them the neck feathers, both dorsally and ventrally,
but especially the former, have a large part of their distal borders
metallic green, as in a cock bird. These neck feathers show distinct
traces of the white ring characteristic of P. torquatus ^. The feathers
of the back are of normal female appearance ; and so are the wing

44 Sterile and Hjihvid Pheasants

feathers, though in three cases both primaries and secondaries show a
slight tendency towards the male pattern. The breast feathers are
intermediate in colour between the cock and hen, being of a rufous
brown colour with some black patches, and occasional feathers showing
the purple and green lustre characteristic of the cock. The tail is of
the female type as a whole, but the black bars on the feathers tend to
be nari'ovver as in the cock.

A partial a.ssumption of cock's plumage by female pheasants, as
above described, appears to be of fairly common occurrence, and it is
invariably associated, as far as the evidence goes, with some abnormality
of the ovar^'. Examples of the converse event, namely, the assumption
of hen's plumage by the cock, is far rarer, though a few cases have been
recorded (Hammond Smith, The Field, Feb. 25th, 1911, p. 384), and in
these cases the testes of the male show no abnormality or signs of
degeneration. In reply to enquiries Mr Hammond Smith writes to
me — "My experience is that female birds frequently assume male
characters as to plumage, and that this change is as a rule associated
with atrophy of the ovary which is often represented by merely a small
patch of black pigment. But I have never seen any trace of male
internal organs in these birds. This is a very common thing to see,
viz., the female assuming the male secondary sexual character. I have
seen it in fowls, ducks, grouse and pheasants. My special interest is
and has been for years to obtain specimens of the opposite change,
namely cocks assuming female characters. This is far more rare, in
fact I can only find three instances, one in the Ibis, 1897, one in Tring
Mu.seum, and the partial change mentioned in Shattock and Selig-
mann's paper {Trans. Path. Soc. London, Vol. Lvn. 1906). For eight
years I have looked for such a bird, and my friends have also done the
same; this winter I have got three specimens. Curiously although in
cases of female allopterotism the ovary is abnormal, in these cases of
male allopterotism the testes appear to be normal ; in each of my three
cases the testes have been carefully examined by S. G. Shattock of the
College of Surgeons."

A. Bi'andt (6), writing in 1889, refers to " Arrhenoidie," or the
assumption of male plumage by the female, as a common occurrence in
various birds, and cites instances among Gallinacei, Passeres, Scansores,
Grallatores and Natatores. He remarks also that these cases are
always accompanied by ovarian abnormality. The opposite phenomenon,
" Thelyidie, " or the assumption of female plumage by the male, is, ac-
cording to Brandt, much rarer, but he is not so clear in regard to these

GrEOFFREY 8mITH AND MrS HaICt ThOMAS 45

cases, as he cites as instances some birds with hermaphrodite internal
organs which obviously come into a totally different category. On the
whole, however, Brandt's evidence agrees essentially with Hammond
Smith's and our own. Mrs Haig Thomas on the other hand has in her
records a case of a male common j)heasant, probably about eight months
old, with partial female plumage, in which the testes were exceedingly
small, about the size of a six weeks old male chick. It seems to me
probable that this was a case of a late-maturing cock bird, in which
both male plumage and normal testes would have developed later. My
reason for suggesting this is that I have at present a breed of white
Leghorn fowls in which the cock birds fall into two classes, those which
assume full male plumage, spurs and comb, and which become internally
mature about six months after hatching, and those which it is impos-
sible to distinguish from hens for eight months after hatching and
which then gradually assume the male characters. The two characters
of early-maturing and late-maturing are hereditary characters which
appear to segregate in inheritance, but the exact nature of this here-
ditary transmis,sion has not yet been worked out.

Now if the above arguments are substantiated, it would seem that
the appearance of the secondary sexual characters of the opposite sex
in the female and in the male bird may be due to two different cau.ses ;
in the female it appears to be due to atrophy or abnormality of the
ovary, and in the male to a transference of the female plumage inde-
pendently of any abnormality of the testes. The case of the female is,
therefore, one of correlated differentiation, where the atrophy of the
ovary liberates some stimulus for the development of the male charac-
ters, while the case of the male is due to hereditary transference, wheie
a particular set of female secondary sexual characters are transferred
to the male without the structure or function of the testes being
affected in any way.

That such hereditary transferences of secondary sexual characters
from one sex to the other often occur, without the normal reproductive
capacity of the individual being affected, is well known: common
instances are the hen-plumage of the Sebright Bantam cocks, and the
appearance of spurs in the females of many breeds of fowls. In both
these cases the males and females are perfectly normal reproductive
individuals ; all that has happened to them is the transference in
heredity of a particular group of secondary sexual characters which
properly belong to the opposite sex.

We must, therefore, clearly distinguish between two entirely

46 Sterile and Hybrid Pheasants

distinct methods by which an individual can assume secondary sexual
characters of the opposite sex, (1) by correlated differentiation, where
the abnormality of the gonad induces the developement of characters
proper to the other sex, (2) by hereditary transference, where the
secondary sexual characters are abnormally distributed in heredity
without the gonad being affected at all.

It is on the basis of these two main subdivisions that a logical
classification of secondary sexual characters, both normal and abnormal,
should be founded, but it is not the intention in the present paper to
apply the principles of this classification any further than to the facts
under consideration. It is clear, at any rate, that the four female
"mule" pheasants described above are instances of the first phenomenon,
viz., correlated differentiation. Three other instances of this process
are afforded by the hj'brid females, M, B and G, and the sterile pure-
bred female, K. The last-named bird, K, was a pure-bred Formosan
variety of the Chinese pheasant, P. torquatus, hatched by Mrs H. T.
May from stock obtained from Mr W. Trumperant Potts of Correen
Castle, Ballinasloe. This bird was hatched in May 1909, and in Sep-
tember of that year was observed to " feather up " like a cock, and
showed rather more of the red face-skin than a normal female. During
this year it also developed a few feathers of the male colour on the
inner part of the thigh. In July 1910 it began to tread the hens
placed with it and was heard to utter the male cry, though no male
plumage appeared until October when a few dark feathers were noticed
below the bill. These were lost however at the moult. In 1911 the
bird again developed male behaviour and at the September moult
began to assume male plumage to a marked degree. During October
it acquired a brown-tinted flush over the body, wings and tail, dark
feathers appeared on the breast, the white collar characteristic of the
male of this species appeared, and the throat and neck feathei-s were of
the lustrous green colour found in the male. It was dissected in the
summer of 1912, when it showed these characters, and it was found
that the ovary was a shrivelled body about six millimetres across,
consisting histologically of interstitial and stroma cells with a large
invasion of fibrous tissue, but no trace of oocytes at all. The oviduct
was in a remarkable condition, being greatly inflamed and hyper-
trophied, although the bird had never laid.

The other three sterile females were all hybrids. 3f was a hybrid
between Reeve's $ and versicolor J*. It never laid eggs, and on dis-
section during the breeding season at three years old, the ovary was

Gbob^frey Smith and Mrs Haig TiioiMAs 47

found to be small, though containing some small oocytes ; the oviduct
was small and comparatively undeveloped. This bird had practically
typical female plumage, though the pattern of some of the feathers,
e.g. the interscapulars, showed a tendency towards the Reeve's male.

Of the two hybrids between Reeve's $ and Formosan ^ , B had
again typically female plumage with a trace of male pattern in some
feathers. The ovary of this bird, at two years old and in the breeding
season, was markedly degenerate, there being no oocytes but only inter-
stitial and stroma cells. The genital ducts were peculiar in that there
were two oviducts (an abnormality of occasional occurrence in Galli-
naceous birds), which showed the same inflammation and hypertrophy
as in the sterile female K. The other hybrid G exhibited a strong
assumption of male plumage, green and purple feathers appearing on the
back, neck and breast. The ovary in this bird was highly degenerate,
consisting of a few minute nodules without any trace of oocytes, and
the oviduct was exceedingly small and undeveloped.

Taking all these sterile females together we must observe a
considerable variation in the degree to which the secondary sexual
characters are altered, and in the degree to which these alterations are
correlated with the condition of the gonad.

The females A, G, D and E showed absence of ovary associated
with very small oviduct and a marked assumption of male plumage.
M had a small ovary with oocytes, a small oviduct and very slight
assumption of male plumage. B had a degenerate ovary without
oocytes, hypertrophied oviducts and very slight assumption of male
plumage. K had a degenerate ovary without oocytes, hypertrophied
oviducts, and a strong assumption of male plumage. G had a degene-
rate ovary without oocytes, a small oviduct and a strong assumption
of male characters. We see therefore that in a female with ovaries
degenerate in a marked degree we may find either a small rudimentary
oviduct or a greatly hypertrophied one, and we may find a very slight
or a highly marked assumption of male plumage, without being able to
trace any correspondence in the degree of degeneration of the ovary.
Nevertheless the evidence is ample to show that degeneration of the
ovary is accompanied by some degree of assumption of male plumage
in every case, and by a marked assumption in the majority of cases,
and we are warranted in concluding that there is an organic correlation
between plumage and ovary in the case of pheasants of such a kind
that the presence of a normal functional ovary is necessary for the full
developement of the female plumage in a female pheasant.

48 Sterile and Hybrkl Pheasants

There is ;i certain amount of experimental evidence on other birds
which confirms this result. Thus Guthrie (4) records the case of a hen
which assumed cock's plumage after the removal of the ovary, and
Goodale (5) observed the developement of Drake's plumage in some
Ducks from which he had removed the ovaries.

We must suppose that the ovary in a female bird normally exerts
an influence on the developement of the typical female plumage, but
we are not in a position to say how this influence is exerted, whether
by means of a specific hormone, or by entering into a chain of metabolic
processes and stimulating the production in excess of substances which
are formed elsewhere. In previous studies the latter mode of action of
the gonad upon the body has been advocated in opposition to the ordi-
nary hormone theory, and it may be suggested that in the cases before
us the wide range of variation in the degree to which the secondary
.sexual characters are altered when the gonad shows no corresponding
variations in its degree of degeneracy, points to other organs of the
body, in addition to the gonad, taking part in the coiTelation.

It will be noticed that these cases of assumption of male characters
by the female as the result of ovarian degeneracy are the converse of
the case of the crabs infected by Sacculina where the males under the
influence of the parasite assume female characters.

The two cases are not, however, completely parallel, because in the
birds we are dealing with a mere suppression of the ovary, while in the
crabs it is not merely a suppression of the testis but the active feminis-
ing of the Sacculina which brings about the changes.

Nevertheless the two cases are parallel, in so far as they prove that
certain male secondary sexual characters are latent in the body of the
female bird which can be made patent by the removal of the influence
of the ovary, and that certain female secondary sexual characters are
latent in the body of the male crab which can be called forth by appro-
priate stimuli.

It would be possible to use these facts in favour of that particular
Mendelian theory of sex which looks upon one sex as heterozygous for
sex and the other homozygous, and to urge that in Birds the female
sex is heterozygous, the male homozygous, and in certain Crustacea the
male is heterozygous and the female homozygous. It is certainly a
remarkable fact that on the one hand in Birds the assumption of male
characters by the female is frequently associated with abnormality of
the ovary, while the assumption of female characters by the male is
independent of any reproductive abnormality but is due to hereditary

GrEOFFREY BmITH AND MrS HaIG ThOJIAS 49

transference ; in Crustacea on the other hand the converse is true, viz.
that assumption of female characters by the male is generally associated
with abnormality of the gonad, while the assumption of male characters
by the female is unaccompanied by abnormality of the gonad.

This might be taken to indicate that in Birds the female carries
the male primary se.xual quality in a latent state which on the removal
or abnormality of the ovary comes into activity and stimulates the
developement of the male secondary sexual characters, while in
Crustacea the male carries the female primary sexual quality which on
removal or abnormality of the testis or owing to the activity of some
special stimulus comes into activity and stimulates the developement
of the female secondary sexual characters.

Although it would be rash to press this argument, the apparent
difference in the reaction of the sexes in Crustacea and Birds would
seem to argue a different physiological constitution in respect to the
distribution of the primary sexual character to the two sexes. In
regard to the distribution of the secondary sexual characters to the
two sexes, it appears probable that each sex is carrying the opposite
secondary sexual characters in a latent state. This is most clearly
proved by Mrs Haig Thomas' hybridising experiments with pheasants
where she has been able to prove that the female can transmit to her
male offspring the male secondary sexual characters of her species, and
also that the male can transmit to his female offspring the female
secondary sexual characters of his species. This fact is well illustrated
by the hybrid pheasants dealt with in this paper. The detailed
evidence, however, cannot be given here but will be described in
a separate paper by Mrs Haig Thomas.

3. The secc-ratiu of hi/brid pheasants.

In an interesting paper Guyer (2) has called attention to the
preponderance of males over females produced by crossing different
varieties among Gallinaceous and other birds. In Part 9 of my
studies (3) I alluded to this observation and recorded that as far as my
experience went the same preponderance was met with. The following
statistics which Mrs Haig Thomas has derived from her extensive
experiments in raising hybrid pheasants are of exceptional value,
because the sexes of the birds were determined in doubtful cases by
dissection, and also a record was kept of the number of infertile eggs,
of the number that died in the shell, and in certain cases the sex of the
birds that died in the shell was determined. They also include a very

Joum. of Gen. iii 4

50 Sterile and Hybrid PheasanLs

large number of cases, so that the irregularities of chtuioe are ruled
out.

The first numbers that will be given refer to the sterile hybrids
raised by crossing Reeve's $ with males of F. torquatas, var. fornio-
saiius, and P. verniculor. The numbers arc as follows: —

Infertile and Dead in

Total eggs Addled sliell Sex Totals

99 68 1 6? 16cr

The sex-ratio is here 6 $ to 16 (/" or 266 per cent, of males. The
especially interesting point brought out by these figures is the fact
that the number of birds which died in the shell is only one, so that
the disproportion in the sexes cannot be accounted for by supposing
that more female birds died in the shell than males. The cause of the
disproportion must be sought for at an earlier stage, either at so early
a stage of developement that no trace of a developing chick could be
detected in the egg, or else at fertilisation itself. In other words the
disprojjortion of the sexes appears to be due not to a selective mortality
of the sexes, but to something tantamount to selective fertilisation.

We may now give the total figures relating to all the cross-bred
pheasants raised by Mrs Haig Thomas. It must be noted that these
cross-bred pheasants were not sterile hybrids in the strict sense of the
term though they exhibit a considerable degree of sterility. The
crosses were mostly between Silver and Swinhoe Pheasants, and Golden
and Amherst, and Forniosan and versicolor.

Table of Sex-ratios in Hybrid Pheasants.

Year

Total eggs

Infertile and
Addled

Dead in
shell

Sex Totals

1909

544

3.56

17

2? Os

55?

83 cf

1910

281

2.32

12
1? 3j

17?

19 c?

1911
1912

424
273

306
117

11

1 ? -2^
12

27?
36?

63 <?
63 J

Totals

1522

1011

52

4? 5<f

135?

228 J

Note. The death-rate of hatched birds from 1909 — 1912 was 143 males to 55 females
thus showing a preponderance of deaths among males. 96 out of the total 1522 eggs were
not incubated.

The sex-ratio is here 135 $ to 228 f^ or 168 per cent, of males.
This percentage represents a considerable drop from that obtained from

Geoffrey Smith and Mrs Haig Thoimas 51

the perfectly sterile hybrids, but nevertheless the preponderance of
males is striking enough.

Now by referring to the column Dead in shell, it will be seen that
52 chicks died in the shell, and on the sujjposition that all these were
females it would bring the total number of females up to 191, which is
still considerably below the total number of males, viz. 238. But in a
few cases the sex of the birds dead in the shell was determined and in
four cases the sex was female and in five male. There is therefore no
reason for supposing that the 52 chicks dead in the shell were all females;
on the contrary as far as the few figures go there is a slight preponder-
ance of males. Taking this result in conjunction with the result
obtained from the first series of figures, it is clear that a selective death-
rate during developeinent, at any rate after the rudiment of the chick
is visible, does not occur. The cause of the disproportion of the sexes
must be sought in the 1011 eggs out of 1426 which were either not
fertilised or else ceased to develope at so early a stage that no trace of
a blastoderm or chick could be detected. If we accept the theory which
at the present time has the largest number of supporters, that sex is
determined at fertilisation by the nature of the gametes that make up
the zygote, we must suppose that in these cases of hybridism successful
fertilisation occurs most frequently between gametes which will form a
male zygote than between those which will form a female. This is
equivalent to selective fertilisation, and the facts recorded certainly
constitute very strong evidence in favour of this process occurring, a
process which has often been suggested as a possibility though never
directly proved.

Summary.

1. Evidence is brought forward to show that the sterility of the
male and female hybrid pheasants is due to abnormalities occurring at
the synapsis stage of the reduction division.

2. The spermatozoa of the hybrids are either deformed, with
chromatic beads and thickenings on them, or else of abnormal size,
double-sized spermatozoa being of frequent occurrence as in hybrid
pigeons. The oocytes in the hybrid females fail to grow owing, probably,
to the early synapsis which should take place in the 19 day old embryo
having gone wrong.

3. Partial assumption of cock's plumage by the female as the
result of ovarian degeneration is shown to be of fairly common occur-
rence. The oviduct in these cases may be either atrophied or greatly

4—2

o2 Sterile (unf Hi/bn'tf P/ieasants

hypertrophicd. Assuiiiptiuu ut hen's plumage by the cock bird is much
rarer and is generally due to hereditary transference of this character
independently of any disturbance of the testes or the reproductive
function.

4. The sex-ratio among hybrid pheasants gives a large preponder-
ance of males over females (228 J' to 135 ^ ). This preponderance
cannot be accounted for by a greater death-rate of female chicks in the
shell or subsequently, and must be presumably due to a selective ferti-
lisation by which more male zygotes are produced than female, or else
to a selective death-rate at so early a stage of developement that no
trace of an embryo could be detected.

EXPLANATION OF PLATE I.

[All the figures are drawn with the camera under a 4 eyepiece and ,'.j objective.
Magnification about 1270.]

Fig. 1. Six spermatozoa of normal common pheasant. {P. torqiiatus.)

Fig. 2. Specimens of spermatozoa of hybrid Reeve's ? by versicolor s ■ They show

abnormality of size, and deformity of shape in some cases.
Fig. 3. Section of testis tubule of hybrid Eeeve's ? by rerslcolor i . Spg. spermatogonia.

Spa. 1, primary spennatocytes showing abnormal spireme stage. Sp. secondary

spermatocytes, spermatids and spermatozoa all showing a high degree of abnormality

and degeneration.
Fig. 4. Section through an outer part of degenerate ovary of hybrid Reeve's ? by Formo-

san i (hybrid £). Ep. external epithelium. (. a group of interstitial cells.
Fig. 5. Section through an inner part of degenerate ovary of B. i. group of interstitial

cells. /. fibrous tissue, ci/. a cyst.

LIST OF LITERATURE.

1. Heinkich Poll. " Mi«chliiig«tudien, vi." Archiv f. Mih-oslcopische Aiiatomie,

Bd. 78, 1911.

2. GnYER, M. F. " On the Sox (if Hybrid Birds," Biological Bulletin, Vol. .\vl

1909, p. 193.

3. Smith, G. " Studies in the Experimental Analy.si.s of Sex. Part 9. On

Spermatogenesis and the Formation of Giant Spermatozoa in Hybrid Pigeons."
q. J. M. S. Vol. Lvni. 1912.

4. Gdthbie, C. G. Journal of Expcriiiuintal Medicine, Vol. xn. p. 269, 1910.
.5. GooDALE, H. Biological Bulletin, Vol. .\x. 1910.

6. Brandt, A. Zeitschrift f. Wiss. Zool. Vol. xlvhl p. 101, 1889

JOURNAL OF GENETICS, V0LIII.N5 |.

PLATE

ly . ^j^i

^^i '^^ %

3.

M^

c*^

I-
I

fT^
.^l

^(^^

E . Wilson , Cambridge,

PRELIMINARY NOTE ON HETEROSTYLISM
IN OX ALLS AND LYTIIRUM.

By N. barlow.

The following paper deals with the heredity of the three forms in
trimorphic heterostyled plants. The work is fxr from complete, bnt
is perhaps worth publishing as the numbers obtained are large, and
certain definite conclusions have already been reached. No factorial
interpretation of the results, however, has at present been devi.sed.

Both Darwin (1) and Hildebrand (2) worked at the descent of the
three forms in trimorphic species, but in Darwin's case only as a side
issue. Moreover their numbers are not in agreement, and, with the
exception of Darwin's self-fertilized Lythrums, the results are no great
help to us at present.

Bateson and Gregory (3) have shown that dimorphism in Primula
is a case of simple Mendelian inheritance. The long-styled plants
are always recessives (RR), whereas the short-styled plants are either
homozygous for the dominant short style (BD), or heterozygous, con-
taining both D and R. The unions therefore between long- and
short-styled plants can be of two kinds ; the F^ will either consist
of short-styled plants only, or there will be equality of longs and
shorts.

In dealing with trimorphism, there is apparently a much greater
degree of complexity than might be expected in the light of these
simple results obtained for dimorphic species.

The three forms within each trimorphic species only differ from
one another in the developement of their sexual organs, and in the
size and sometimes in the colour of the male sexual cells themselves.
Each individual has two distinct sets of male organs, and one female
oi-gan, arranged in three tiers. Thus each of the females of the
three different types can be pollinated by six perfectly distinct males,

54

Heterostylism in Oxalis

making eighteen possible pollinations within the species, of which six
are self- and twelve cross-fertilizations. The six self-fertilizations are
always partially or quite sterile. The six " illegitimate " cross-fertiliza-
tions ( ? and ^ of different tiers) are likewise infertile, and only the
six "legitimate" unions are fully fertile. This partial sterility is a
great obstacle in determining the gametic constitution of individuals,
as selfed seed is difficult to obtain in any quantity and difficult to
raise ■ nor can the " illegitimate " unions be relied on to produce
sufficient seed.

Long_x_Short

on X LoO<i

Fig. 1. Showing the aiiaiigeiiient of tlie styles and stamens in Long-,
Mid-, and Short-styled plants. The connecting dotted lines
indicate the six "legitimate'' forms of nniou.

Sterility must prove to be an integral part of the problem when
it is fully worked out. The relation to the main issue, however, still
remains obscure.

It is significant that the three forms have different average
degrees of fertility even when producing their fullest complement of
seed. The relative numbers of seeds in Lythrum salicaria can be
given as follows: Long-styled 100; Mid-styled 130; Short-styled 81.
(Darwin, Forms of Flowers, p. 143.) Also the different degrees of
sterility in different self-fertilizations and "illegitimate unions "seem
constant. Again, the mid-styled form is more productive than the
other two; but not ecjually so tVtr all illegitimate unions. Thus Darwin
got as many as 69 5 seeds per capsule in the cro-ss Mid-styled x long
stamens of Short-styled, but on the other hand not a single seed for
Mid X own from short stamens.

K Barlow 55

The fact that the two sets of pollen on one plant behave differently
towards the same female shows a segregation in the individual of the
factor (or absence of factor) causing incompatibility'. This constitutes a
marked difference between heterostj-led plants and such self-sterile
species as Linaria vulgans (de Vries (4)) and Cardamine (Correns (5)).

Oxalis valdiviana was selected for experiment, owing to its annual
habit and its long flowering period. This plant has a very high
degree of self-sterility, and it was found difficult to proceed with it
alone. Lythrum salicaria has been found to be more self-fertile
(C. D., Forms of Flowers) ; also the normal capsules set over 80 seeds,
whereas in Oxalis there are onl}' from 10 to 1.5. Experiments were
started on this species in 1911 so that we have as yet only the F^.

The plants were obtained from the Cambridge Botanic Gardens, and
from wild seed. Two plants of the variety rosea, growing in the Chelsea
Physic Garden, were also used.

Oxalis valdiviana.

Procedure.

In 1907 four long-, four mid-, and four short-styled plants were
chosen from self-sown garden plants. All families subsequently raised
are derived from these twelve original parents.

During the summers of 1907, 1908 and 1909 the plants were kept
under wire cages covered with muslin ; or each flower-head was sepa-
rately covered with a paper bag. No crosses were made in 1910. In
1911 and in 1912 the plants used were either isolated in bee-proof
greenhouse partitions or covered with paper bags. Bees visit this
plant very freely. It was sometimes difficult to frustrate their per-
sistent attempts to get at flowers temporarily uncovered for artificial
fertilization. Insect visits may be a cause of irregularity in the
results, and will be referred to later, with other possible causes of
error.

Owing to the violent expulsion of ripe seed, each capsule has to be
separately covered with fine muslin before it reaches maturity.

The immature anthers were removed from the females used for
crossing in 1907. In subsequent years castration was not thought

' A flower of the short-styled Oxalis valdiviana was observed to have one of the mid
length tier of stamens at the level of the long stamens. The pollen from this aberrant
stamen was carefully tested on a long- and on a mid-styled plant, and was found to behave
exactly as "long" pollen. The length factor of the stamen is bound up with the inhibitory
effect of the pollen ; the abnormal growth of this stamen had brought with it the inhibition
of its pollen for a mid-styled plant.

56

HeterostjfUsm in Oxab's

necessary, owing to the degree of self-sterility. The risk incurred was
slight, and will be referred to later. In mid and short females, however,
the tiers of ripe anthers above the level of the stigma were always
carefully removed before the foreign pollen was administered. In the
controls for this procedure no seed set. In male parents pollen at
a higher level than that used was removed in a similar manner.

Unless otherwise stated, the cross denoted by "Long x Mid,"
" Mid X Short," etc., is the fertile cross between the two forms, that
is, Long-styled % x poUen from long stamens of Mid-styled ; or Mid-
styled $ x pollen from mid stamens of Short-styled, etc.

Results.

Four generations have been raised. The results of the crosses
Long X Mid, and Long x Short, in the four years, together with
their reciprocals, are given in Table I and Table II.

TABLE I.

.

0.

mlia vafdiriana.

Long-styled

female X

Mid-styled male

Mid-styled female xLong-

styled m

Longs

Mids

Shorts

Lougs

Mids

Shorts

1908

6

2

0

5

4

0

1909

78

C8

0

50

52

4

1910

31

4.5

0

140

137

2

1912

23

22

0

14

12

0

Totals 138 137 0

Reciprocals added together:—

209

205

Longs Mids

Shorts

138 137

0

209 205

6

Totals 347 342

0

TABLE n.

Oxalis valdiviana.

Long-styled female x Short-styled male

Short-styled female x Long-styled male

Longs Mids Shorts

Longs

Mids

Shorts

1908

10 0 9

28

3

22

1909

50 3 38

52

6

08

1910

138 0 135

03

0

58

1912

0 0 9

8

0

8

Totals

203

191

151

15G

N.

Barlow

;ether :

—

Longs

Jlids

Shorts

203

3

191

151

9

156

57

Totals 354 12 347

A long crossed with the legitimate pollen of a mid, or reciprocally,
gives equality of the parental forms, with the occasional presence of
the third non-parental type. Amongst a progeny of 695, the short-
styled form occurs 6 times, or 1 per 117. Whether this is due to
possible causes of error, or has a real significance, I do not at present
feel qualified to say, though I incline to the latter opinion.

Again, a long crossed legitimately with a short, and i-eciprocally,
gives a predominance of the parental forms, with the mid-styled
plant occurring 12 times in a total of 713, or 1 per 59.

The crosses between mid-styled plants and short-styled plants give
an entirely different result. No regularit}' is visible if the numbers
are given in an unanalysed tabular form.

TABLE 111.

Mid-styled female x Short-styled male Short-styled female x Mid-styled male

Longs Mids Shorts Longs Mids Shorts

1908

3

4

3

4

2

12

1909

17

10

28

16

16

24

1910

44

100

174

103

13G

24G

1912

82

241

341

95

241

27G

Totals 146 355 546 218 395 558

Analysis of F., Mid x Short and Short x Mid, flowering 1910.

In Table IV, the first group is the addition of all cases where there
is at least one Long-styled plant amongst the four grandparents. We
get 118 Longs : 121 Mids : 270 Shorts — a possible approximation to
the ratio 1:1:2.

In the second and third groups there is complete absence of grand-
parental Longs. The second gives 23 : 23 : 49 — probably the 1:1:2
ratio. But the third gToup gives Mids and Shorts equally, with an
occasional Long, the numbers being 6 : 102 : 101.

Clearly all Mids, or all Shorts, are not similarly constituted. The
above families do not give clear evidence as to which form bi'ings in
the difference. There are certainly two, and probably more than two,

58

Heterostylism in OxaUs

different types of Mid-styled plant, and possibly two different types
of Short-styled plant. We get clear evidence for different Mids in
the 1912 numbers to follow. But in Table IV, if we consider cross 31
and its reciprocal 32, we get 65 Longs : 59 Mids : 130 Shorts. In
cross 33 and its reciprocal 34, where the same individual Mid parent
is used, we get 19 Longs : IS Mids : 87 Shorts. This may be taken as
doubtfid evidence for a second type of Short-styled plant.

TABLE IV.

0. valdiviana F.,.

Tlic letters in brackets show the extraction of the parents.

Cross Number

Parents

Longs

31
33
32
34
37
70
71
72
75

Mid (L X M)
Mid (L X iU)
Short (L X S)
Short (Sy.L)
Short (L X S)
Short (S X L)
Short (lUxS)
Short (])/ X S)
Short (.U X S)

X Short (LxS)
X Short (.S' X L)
X Mid (L X M)
X Mid (L X M)
X Mid (M X L)
X Mid (L X M)
X Mid (Z, X iU)
X Mid (L X M)
X Mid (L X M)

28

11

37

8

12

19

3

0

0

23

8

36

10

13

20

0

3

52
50
78
37
14
25

0

2

12 /

41

c Mid(.l/x.S')

Totals

118

121

270

X Short (il/ X S)

2

4

6

1

c Short (M X .S)

X Mid {M X S)

17

16

37

15

and 10

Short (.1/ X S)

X Mid (M X S)

4

3

6

Totals

23

23

49

9

Mid (M X ,S')

X Short (M X ,S')

2

28

29

28

Mid (M X S)

X Short (,1/ X .S')

1

i;i

IS

44

Mid (il/ X S)

X Short (M X .S')

0

22

It;

17

Mid (M X S)

X Short (il/ X .S')

0

6

3

7

Short (M X H)

X Mid (il/ X S)

1

7

7

8

and 11

Short (.1/ X S)

X Mid (A/ X S)

2

20

28

Totals

6

102

101

III

The small letters show reciprocal crosses.

Analysis of F^ Mid x Short and Short x Mid, flowering 1912.

Unfortunately the F-^ parents fin- the F^ generation wx're chance
survivors through the winter of 1910. There remained in 1911
three Short-styled plants and seven JI id-styled plants to carr}- on

N. liARLOW 59

the race, besides some Longs. Moreover these ten plants were not
all in the same place, and certain crosses, which might have been
elucidating, were thus precluded.

TABLE V.

0. valdiviana F^.

Parents

Cross Number -' Longs Mids Shorts

Mid ? Short S

10 9/2 (MxS) X 9/1 (MxS) 0 30 34

11 9/3 (il/xS) X 9/1 (MxS) 0 30 24

12 9/2 (i¥xS) X 70/5 (MkS) 0 24 23

13 9/3 (MxS) X 70/5 (UxS) 0 40 32

6

Totals ... 0 124 113 =A

16 70/2 (ilfxS) X 70/1 (MxS) 1 17 34

17 70/3 (MxS) X 70/1 (il/x,S') 1 27 3G

18 70/4 6 (il/xS) X 70/1 (M x S) 0 G 17

Totals ... 2 50 87

14 70/4 (il/xS) X 9/1 (IfxS) 8 8 8

19 70/4 (il/xS) X 70/5 \m-aS) 43 30 72

Totals ... 51 38 80

20 37/1 (MxS) X 9/1 (MxS) 10 17 29

15 37/1 (MxS) X 70/5 (Jl/x,5) 17 11 29

Totals ... 27 28 58

Short ? Mid i

1 9/1 (MxS) X 9/2 (MxS) 0 28 25

2 9/1 (ilixS) X 9/3 (MxS) 0 4 4

3 70/5 (MxS) X 913 (MxS) 0 43 39

4 70/5 (MxS) X 9/2 (MxS) 0 33 21

Totals

0

108

89

= A'

70/1 (MxS) X 70/2 (MxS)
70/1 (MxS) X 70/3 (MxS)
70/1 (MxS) X 70/16 (MxS)

3
0
0

18

2G

9

15

26

3

Totals

3

53

44

=B'

70/5 (MxS) X 70/4 (MxS)

84

76

133

= C'

5 70/5 (M X S) X 37/1 (M x S) 8 4 10

A and .4' = reciprocals.
B and B' =
etc.

00 ffeterofityUsm in Oxalis

These figures increase our knowledge on certain points.

(1) There is no evidence against the supposition that male and
female gametes of a plant are carrying the same qualities. Compai-e
A and A' etc. In the four groups A — D there is good agreement
with their reciprocals A' — U.

Note, however, the excess of Shorts in B, and the excess of Mids
in B'.

(2) Two of the three Shorts used are similar in constitution (9/1
and 70/5). It was not possible to cross the third Short with any of
the Mids used with 9/1 and 70/5, so no comparison can bo made.

(3) The seven Mids used can be classed according to their offspring
when crossed with the same Short.

Four different Mids were used with the Short 9/1; — giving:

0 Longs
0 „
8 „
10 „

28 Mids : 25 Shorts Cross 1 Mid-parent 9/2

30 „ :24 „ „ 11 „ 9/3

8 „ : 8 „ „ 14 „ 70/4

17 „ :29 „ „ 20 „ 37/1

Again these four Mids were crossed with the Short 70/5 ; — giving :
0 Longs : 33 Mids : 21 Shorts Cross 4 Mid-parent 9/2
0 „ :43 „ : 39 „ „ 3 „ 9/3

84 „ : 76 „ : 133 „ „ 6 „ 70/4

8 „ : 4 „ : 10 „ „ 5 „ 37/1

Clearly the Mid-pai-ents 9/2 and 9/3 aie similar, and differ in con-
stitution from 70/4 and 37/1.

In the case of the remaining three Mids used as parents for the
Fi generation, there is no comparative evidence on their conformity
with one or other of the above classes, or on the possibility of their
representing a third type of Mid-styled plant.

TABLE VI.

Showiny the ratios yivini hy tlie Jotir Mid-parents crossed with two
similarhj constituted Shorts; and of tlie three remnininy
Mid-parents crossed with Shorts of unknown constitution.

Mid-Parent

Longs

Mids

Sliorts

9/2

0

115

103

9/3

0

117

99

70/4

135

114

213

37/1

35

32

08

70/2

4

35

49

70/3

1

53

62

70/4 i

0

15

20

N. Barlow

61

Self- Fertilizations and Illegitimate Crosses.

Most of such fertilizations have failed. Over 250 have been
attempted, from which S7 plants have been raised.

Longs Mids Shorts

Long-styled X own mid pollen (3 plants) ... 46 0 0

Long-styled X own mid + short pollen 12 o ^

Totals

58

Mid-styled x own long + short pollen

Mid-styled x long pollen of different Mid
Mid-styled x short pollen of different Mid

Short-styled X own mid pollen

Short-styled x mid pollen of different Short
Short-styled x own mid -I- long pollen . . .

Table VII shows the result of all the self- or illegitimately-fertilized
capsules, together with the germination of the seed.

r^

5

3

■•■ lo

i

0

3

1

0

1

0

0

0

0

4

0

0

2

0

0

3

TABLE VII

Parents

Flowers
fertilized

Capsules
setting

Seeds

Germinations

Giving

Year

Female

ISale

Longs

Mids

Sho

1907 and 9

L

X

mL [6]>

45

11

60 +

46

46

0

0

1907 and 9

L

X

sL [8]

15

0

0

0

—

—

—

1908 and 9

L

X

m + sL [3]

12 4-

44-

64

12

12

0

0

1907

L

X

sM [5]

5

0

0

0

—

—

—

1907

L

X

mS [5]

5

0

0

0

—

—

—

1907 and 9

M

X

IM- [11]

46

4

20

4

3

1

0

1907 and 9

M

X

sM^ [11]

24

1

1

1

1

0

0

1908 and 11

J/

X

l + sM [3]

many

6

16

■ 11
4

3
0

5
4

3
0

1907

.1/

X

sL [5]

5

0

0

0

1907 and 9

S

X

IS [11]

20

0

0

0

—

—

—

1907 and 9

S

X

mS-'' [12]

49 +

4

7

6

0

0

6

1909 and 12

s

X

l + mS\_i]

14-i-man.^

4

7

2

0

0

5

1907

s

X

mL [4]

4

1

3

1

0

0

1

1907

s

X

IM [5]

5

0

0

0

—

—

—

1 All the Mids, two of the Longs, and one of the Shorts which successfully produced
seed, had a common Mid ancestor which seemed to be more capable of producing selfed
seed than the others. See also Table VIII where the descent is from the same Mid-styled
plant. The numbers in brackets after the parentage show the number of ? parents used.

'^ The pollen from another individual of the same type.

62 HeterostijUxin iu Oxalis

Procedure and Pusnible Suurcea of Error.

As has been already stated, the Howers were not castrated before
use, excejjt in the year 1907. The danger incurred can be gathered
from the following facts.

During the five summers of 1907, 1908, 1909, 1911 and 1912 the
total number of capsules setting on covered but untouched flowers
amounted to 14, of which none were on Long-styled plants, eleven
on Mid-styled plants, and three on Short-styled plants. Such capsules
may either arise through some insect introducing foreign pollen or
from a true spontaneous self-fertilization. Also, aphis were present on
some of the plants. But these insects would probably act as self- and
not as cross-fertilizing agents, owing to their stationary habits.

It seems probable that these fourteen capsules did not arise from
any outside pollen having reached the stigmas, from the fact that they
were all formed on Mid- and Short-styled plants. The protruding
Long-styles would be most subject to the contact of any intruder, but
no capsules have arisen on untouched Long-styled plants.

As will be seen from the details in Table VIII these capsules set
very few seed, and less than half of these germinated. Also it must
be remembered that these fourteen capsules occurred amongst many
hundreds of flowers which ojDened each year under cover but were
not used.

TABLE VIIL

Cajmule); sctlimj seed spontaneously.

Giving

Year

Type of Parent

capsules

of seeds

GermiDations

Longs

Mids

Shorts

1907

Mid (1 plant) i

4

10
^27

2

1

1

0

1908

Mid (1 plant)

4

15

7

8

0

1909

—

0

0

0

—

—

—

1911

Mid (1 plant)

H

8

4

0

4

0

Short (1 plant)

3

6

3

0

0

3

1912

Mid (1 plant)

2

2

—

—

—

—

To return to the rare appearance of the third form amongst the
offspring of a fertile Long x Mid and Long x Short cross. The
question arises : Can these be accounted for by the introduction of
a few of the mother's pollen grains ? Where a Long-styled plant is
mother, this is clearly not the case, as self-fertilized Longs give Longs
only. (See also Lythrum, below.) But three Mids have appeared in

' The Mid-plant of 1907 was an ancestor of all the subsequent individuals setting
spontaneous seed. There may be self-setting strains. See also Table VII.

N. Barlow 63

a Long $ X Short ^ cross. If this is an error, therefore, it must be
due to foreign pollen or seed in the soil.

In crosses with Longs as fathers, and Mid or Short mothers,
there is nothing conclusive to show that the non-parental type has
not arisen from the mother's pollen-grains reaching the stigma. But
in Table VIII the Mid-plants setting seed have not given rise to a
Short, nor have the Short-styled plants given rise to a Mid. But the
numbers are few, and the evidence rather negative than positive.

Referring to Tables I and II, we see that the year 1909 gives the
largest number of these possible errors. In this year I have great
suspicion of the earth used for sowing, as in four separate cases one
or two more plants came up in the pots than the number of seeds
sown warranted '.

But putting aside all the results of 1909, we are still left with one
case in 1908 and another in 1910. Whether these rare appearances
must be regarded as accidental introductions, or as a vital part of the
problem, must still remain in doubt.

Lythrum salicaria.
Only the F-^ has so far been raised from this plant.

TABLE IX.

Long-styled female x Mid-styled male Mid-styled female x Long-styled male

Longs Mids Shorts Longs Mids Shorts

6 5 0 93 84 0

Long-styled female x Short-styled male Short-styled female x Long-styled male
50 0 46 123 0 103

8= VI- T-

jttid-styled female x Short-styled male Short-styled female x Mid-styled male

53 63 145 71 56 162

Longs and Mids crossed together give only the two parental forms
in a total of 188.

' None of these four cases coincided with a family in which the non-parental form
made its appearance.

- This cross is under suspicion. The flower spikes were covered either by paper bags
or by muslin. The muslin was found to be an insufficient protection, as several times bees
were seen crawling on the surface, whilst the flowers inside were so near that it might have
been possible for them to receive foreign pollen from part of the bee's body. The above
case was from a muslin covered spike. All plants selected for further generations were
from paper-covered spikes.

64 Heterostylism in Oxalis

Longs by Shorts or Shorts by Longs, with the exception of the one
case already noted, give only Longs and Shorts in a total offspring
of 322.

Mids by Shorts and Shorts by Mids throw all three forms in
numbers not explicable at present.

There is no case of the third non-parental type occurring occa-
sionally as in Oxalis.

The self-fertilizations have mostly failed, chietly owing to the
lateness of the season when they were attempted.

Parents

Female Male Longs Mids Shorts

.1/ X own l + x 0 3 0

.S' X own / + )» 1 1 8

Darwin's numbers for self- and illegitimate-fertilizations are added.

Parents

Female

Male

Longs

Mills

Shorts

L X

III + sL

8

0

0

L X

own III + s

48

0

0

M X

own l + s

1

3

0

il X

fL

17

8

0

M X

IS

14

8

18

S X

own / + III

1

0

8

.S X

iiiL

4

0

8

There seems little doubt that in Lythrum and in Oxalis, as in the
dimorphic Primula, the Long-styled form is the recessive, and can only
give Longs.

SiDiiiiutri/.

1. Reciprocals have always given like results. That is, there is no
evidence that the J' aud $ gametes carry different characters.

2. Long-styled plants selfed give Longs only. Long is the pure
recessive, as in Primula.

3. There are certainly two differently constituted Mid-styled
Oxalis, giving different ratios with the same Short. One gives no
Longs at all, and the other gives Longs in equality with the Mids.
Self-fertilizations can give Longs as well as Mids, and once in Oxalis
all three forms have appeared. There is no comparative evidence
on the result of self-fertilizing the two different types of Mid-styled
plant.

N. Barlow (35

4. There is doubtful evidence of a second type of .Short-styled
plant, which gives a ratio approximating to 1 Long : 1 Mid : 8 Shorts ;
whereas the usual ratio with the same individual Mid is fairly near
1 Long : 1 Mid : 2 Shorts. Self-fertilized Shorts in Oxalis have pro-
duced Shorts only, but in Lythrum one of both the other forms
appeared in a family of ten.

5. A further well-established ratio in a J\] x S cross is

1 Long : )( Mid : n Short,

where n is a number between 13 and 34. This indicates the presence
of yet a third type of Mid or Short, but no comparative evidence is
available.

The work was undertaken at Mr Bateson's suggestion, and I wish
to express m}' thanks for his helj:) and encouragement, and for the
facilities which I have enjoyed at the John Innes Horticultural
Institution during the last two years.

REFERENCES.

(1) Darwin, C. Different Forms of Flowers on Plants of tlie same Species.

(2) HiLDEBRAND, F. Bot. Zeitiuig, 1864, p. 2 ; 1871, pp. 416 and 432 ; 1887,

pp. 1, 17, and 33.
Monatsherichte d. Akad. d. Wissen. :u Berlin, 1866, p. 371.
Lehensverhaltnisse der Oxalisarten. Jena, 1884.

(3) Bateson and Gregory. Proceedings of the Royal Society, B. 1905, p. 581.

(4) DE Vries. Species and Varieties, p. 471.

(5) CoRRENS, Festschrift der medizin-natttrw. Gesellsckaft zur 84. Vers, deutscher

Naturforscher und Arzte, 1912.

Journ. of Gen. iii

SUE UNE RACE DE BLE NAIN INFIXABLE.

By PHILIPPE DE VILMORIN.

Le cas que je me propose d'examiner est celui d'un froment {Tri-
ticum sativmii) issu d'une race cultivee et fixee, semblable par tous ses
caracteres a cette race, et infixable au point de vue de la diminution de
la taille. Cette experience a ete poursuivie pendant de nombreuses
annees et, d'ailleurs, le meme ph^nomene s'est presentd dans deux
varietes differentes.

1°. Beseler's Brown Club Head.

Cette variete, comme aussi celle que nous etudierons plus loin, se
fait reniarquer par un epi can-6 et trfes compact.

1905. Parmi une centaine de plantes du type ordinaire qui est de
taille elevee variant suivant les annees de 1 m. 50 a 1 m. 70, j'ai trouve
une seule plante n'ayant que 80 centimetres de hauteur. II a et<^ sem(5
10 grains de cette plante naine dont 7 sont arrives a complet developpe-
ment, la proportion a ete de 6 individus nains pour un gi-and. Une
plante naine a ete conservee.

1907. 10 grains de cette plante donnent 8 nains et 2 grands. Une
plante est conservee. La meme annee j'ai fait, d'autre part, line expe-
rience sur une plus grande echelle semant environ 300 grains provenant
tous de plantes naines de 1906. 271 plantes se sont developpees dont
207 naines et 64 grandes. Parmi les plantes naines, 4 ont ete con-
servees.

1908. Ces 4 plantes ont donne les resultats suivants :

1° 50 naines et 21 grandes
2° 55 „ 21 „

3° 57 „ 27

4" 44 „ 20

Soit au total 206 naines et 89 grandes.

Cette experience a ete abandomiee au profit de la suivante en pre-
sence de la necessite d'operer sur de grandes quantites et de ne pas
multiplier les lots.

68 Sur line Race de Ble Nam Infixahle

2°. Shirno.

A. Famille blanche.

Cette variete qui n'existe pas en dehors de Verrieres j^rovient d'un
croisement fait en 1886. Elle est parfaitement fix^e et elle est, comme
la pr6cedente, caracteris^e par son 6pi court ct carre. Sa taillc nonnale
varie suivant les annees de 1 m. 20 a 1 m. 40.

1902. Sur 675 plantes on remarque une plante naine de 1 metre
qui e.st con.serv6e.

1903. On trouve 147 plantes naines et 80 gi-andes.

1904. Le semis a ete fait avee les grains melanges de plusieurs
plantes naines ou paraissant naines, conservees I'annee pr^c^dente ; ces
gi-ains donnent 177 plantes naines et 116 grandes.

1905. Le semis se fait dans les memes conditions que 1904, c'est-
a-dire avec des plantes en melange. On trouve 85 naines et 196
grandes.

1906. Semis dans les memes conditions : 238 naines et 95 grande.s.

1907. Semis dans les memes conditions: 197 naines et 88 grandes.
Je n'hesite pas a avouer que cette fa^on de proc6der qui consiste h,

ne pas suivre individuellement chacune des plantes choisies, pr^sente de
graves inconv^nients et que les resultats des deux annees 1904 et 1905
seraient tout a fait d^concertants et inexplicables si, par la suite, il
n'avait ete remi^die a cette erreur; nous aurons a y revenir lors de
1 'interpretation de ces resultats.

1908. Dans la recolte de 1907 il a ete choisi 4 plantes qui ont ete
sem6es s6par6ment et ont donn^ les resultats suivants :

233-08 64 naines et 17 grandes

234-08 55 „ 29

235-08 63 „ 19

236-08 61 „ 22

Soit au total 243 naines et 87 grandes.

1909. La descendance de 233-08 est seule suivie, 5 plantes naines
ont 6te conservees et ont donne :

258-09 10 naines et
259-09 8
260-09 17
261-09 23
262-09 28
Soit au total 86 naines et 53 grandes.

5

grandes

3

}j

14

)J

17

)J

14

>*

Philippe de Vilmorin 69

On remarquera que cette annee nous avons opere sur des semis tres
restraints, de sorte qu'il ne faut pas attacher trop d'importance aux
chiffres ci-dessus ; a cette epoque je m'imaginais encore trouver une
plante fixee au point de vue de la taille naine et je ne croyais plus
avoir besoin d'un grand nombre de plantes dans chaque lot. C'est
cette annee la cependant que pour la premiere fois j'ai conserve quel-
ques plantes grandes afin d'etudier leur descendance.

En tout, dans 258-09, 260-09, 261-09 et 262-09 j'ai conserve
16 plantes dont 11 naines et 5 grandes.

1910. Les plantes naines ont continue a donner une descendance
melangee de naines et de grandes dans des proportions sensiblement
constantes donnant au total 333 naines et 134 grandes soit une propor-
tion de 1 a 2"48, elles ont ete cultivees separement, mais je crois inutile
de donner les proportions pour chacune d'entre elles ; les 5 plantes
grandes n'ont donne que des plantes grandes.

Dans la progeniture des 11 plantes naines il a e'te conserve 54
plantes toutes naines (des plantes grandes ont ete suivies dans la
famille rouge).

1911. Les graines de ces 54 plantes naines ont ete semees separe-
ment et ont donne dans I'ensemble 2791 plantes naines et 1246 plantes
grandes, soit une proportion de 1 a 224.

1912. On a seme, toujours par lot separe, les graines de 74 plantes
dont 24 naines et 50 grandes. De plus un certain nombre de plantes
ont ete envoyees a M. Eateson et etudiees par M. Backhouse.

A Verrieres nous avons eu des resultats concordant absolument avec
ceiix des annees precedentes, c'est-a-dire que les 24 plantes naines ont
donne dans leur ensemble 657 naines pour 293 grandes, soit une pro-
portion de 1 a 2'24. De son cote M. Backhouse, examinant la jaroge-
niture des 6 plantes naines, a trouve 314 naines et 131 grandes, soit
une proportion de 1 a 2-39, et a Merton comme a Verrieres, les plantes
gi-andes n'ont donne que des plantes grandes.

B. Famille rouge.

Cette famille est issue de la famille precedente, son origine remonte
a 1905, annee ou le No. 1746 a donne une plante naine a epi rouge.
Cette variation de couleur avait ete deja remarquee mais negligee les
annees precedentes, elle est evidemment due a un croisement accidentel
avec une race a epi rouge, comme il est prouve par I'examen de la
descendance qui se compose de blancs et de rouges en proportions

5—3

70 Siir line Race de Ble Nain Infixahle

ordinaires ; mais, dans ce cas, le caractere dominant rouge est fixable
et a 6te extrait des la deuxieme generation.

1906. On ne recolte que 6 plantes dont 1 a epi blanc. Toutes les

6 sont naines, ce qui n'a rien de surprenant, etant donne le petit
nombre d'individus. On conserve une plante naine semee a part et 5
plantes naines pour semer en melange.

1907. La plante naine semee a part (224-07) donne 9 plantes dont

7 a dpi rouge et 2 a epi blanc ; au point de vue de la taille on trouve
7 naines et 2 grandes.

Les 5 plantes semees en melange (1851-07) donnent 266 plantes
dont 70 a epi blanc; et, au point de vue de la taille, 146 naines et 120
grandes (meme observation que precedemment). II est conserve 4
plantes naines a epi rouge.

1908. Ces 4 plantes sont semees separement avec les resultats
suivants :

237-08 66 naines et 23 grandes

238-08 65 „ 21

239-08 64 „ 17

240-08 65 „ 18

Soit un total de 260 naines et 7!) grandes.

II n'a pas ete fixit de choix dans le 238-08 qui presente une pi-o-
portion de 63 plantes a epi rouge pour 23 a epi blanc. Les 3 autres
lots sont composes uniquement de plantes a epi rouge (dominant fixe);
on conserve 8 plantes dont 7 naines et 1 grande.

1909. Les 7 plantes naines sont semees separement donnant au
total 189 naines pour 89 grandes, soit une proportion de 1 a 2-12.

La plante (263-09) grande ne donne que des plantes grandes.
Dans la descendance des plantes naines il est conserve 15 plantes
dont 11 naines.

1910. Les 11 plantes naines, toujours examinees separement, don-
nent au total 342 plantes naines et 144 grandes, soit une proportion de
1 a 2'38, les plantes grandes ne donnent que des plantes grandes ; on
conserve 55 plantes dont 2 grandes.

1911. Les 53 plantes naines donnent au total 3048 plantes naines
pour 1213 plantes grandes, soit une proportion de 1 a 2-51.

Les deux plantes grandes etudiees cette annee n'ont donne que des
grandes.

Dans la descendance des plantes naines il est conserve 58 plantes
dont 18 naines.

Philippe de Vilmokin 71

1912. Les 18 plantes naiiies ont donne au total 390 plantes naines
pour 160 grandes, soit une proportion de 1 a 2-43. Les 40 plantes
grandes ne donnent que des grandes. II semble done bien evident qua
la suite d'une experience portant sur un aussi grand nombre d'individus,
les plantes grandes sent incapables de reproduire des jjlantes naines.

3°. Discussion des RfouLXAXs.

Des resultats de cette experience il ressort clairement au moins
deux faits : 1° dans cette race naine, les plantes naines ne se rejjro-
duisent jamais d'une fa9on homogene, la descendance des plantes
naines donnant toujours une certaine jDroportion de jilantes grandes;
2° les plantes grandes de la meme origine donnent une progeniture
homogene de plantes grandes et ne donnent jamais de plantes naines.

II s'agit maintenant d'etudier le rapport entre le nombre de
plantes naines et de plantes grandes dans la descendance des plantes
naines.

On remarquera que si Ton fait abstraction de certains resultats
douteux, surtout dans les premieres annees alors que les mensurations
netaient pas faites avec toute la rigueur necessaire, les gxandes sont
aux naines dans un rapport variant de 1/4 a 1/3.

Au premier abord, et si nous n'avions pas un aussi grand nombre
d'observations, il semblerait que les plantes naines se comportent comme
une forme heteroz3'gote donnant 1/4 de plantes grandes homozygotes,
2/4 de plantes naines heterozygotes et 1/4 d'un type homozygote qui
pour une raison quelconque ne serait pas viable; par consequent, dans le
cas present, ce serait I'homozygote dominant qui disparaitrait. On
pent admettre que ce soit un cas analogue a celui des souris jaunes
etudie par Cuenot(l) et ensuite par Miss Durham(2). S'il y avait repul-
sion entre certains gametes la proportion devrait etre de 3 naines a 1
grande, en supposant naturellement que tons les gametes femelles qui
ne peuvent pas etre fecondes par la premiere categorie des gametes
males le soient par la deuxieme, ce qui serait plausible, le nombre de
grains de pollen etant illimite.

II se pourrait aussi que les gametes males et femelles de chaque
categorie puissent s'accoupler dans les conditions normales mais que le
germe provenant de I'union de 2 gametes portant le facteur resulte en
un germe incapable de se developper. Alors a quel moment de la vie,
I'homozygote dominant disparaitrait-il ?

72 Sur u)ie Race de BU Naiti Injixable

Les grains peiivent ne pas se developper et leiir place rester vide
dans I'epi; pour nous en rendre compte nous avons compte les gi-ains
d'un grand nombre d'epis tant sur des plantes grandes que sur des
plantes naines, arrivant a ce resultat qu'il y a une difference dans le
nombre de grains (59 grains par epi chez les grands ; 50 chez les nains).

II se pourrait aussi que les grains se ferment mais soient incapables
de germer. Nous avons done effectue un gi-and nombre de semis en
terrines, apres avoir prealablement comptd le nombre de grains ense-
mences et ensuite le nombre de plantules sorties de terre.

gr. semes

gr. leves

Famille blanche :

plantes naines

1 2G57

2567

)j J)

plantes grandes

1 805

802

Famille rooge :

plantes naines

3351

3317

La proportion de grains ne germant pas semble done plus forte
dans les plantes naines; je ne crois pas cependant qu'il faille voir la un
phenomene constant expliquant la disparition des homozygotes domi-
nants. En effet, les semis avaient ete faits, plante par plante, et dans
la plupart des cas la difference entre le nombre de grains semes et de
grains lev^s ne depasse pas la normale et est la meme que dans toutes
les vari^tes de bles. Dans les chiffres donnes ci-dessus pour les plantes
naines de la famille blanche, une seule plante, le No. 303-12, dont le
grain etait specialement mal constitue, a donne 307 grains leves sur 345
grains semes.

Si nous faisions abstraction de cette plante nous aurions 2260 a
2312, ce qui est une proportion tres normale. J'ajoute que Ton ne
s'est pas contente de constater la germination des grains, mais que
Ton a conserve les jeunes plantes jusqu'a 1 age on elles sont en etat
d'etre rejDiquees et qu'aucune n'a peri dans cet intervalle.

Nous a%'ions pense aussi qu'une certaine proportion de plantes
pouvait disparaitre dans la suite du developpement, c'est-a-dire entre
le repiquage et la maturite ; mais il n'en est rien et, si forcement
un certain nombre de plantes disparait au cours de la vegetation, la
proportion n'en est pas plus elevee dans les naines que dans les
grandes ou que dans une autre variete de bids.

Comme je I'ai dit, un certain doute sur les resultats pent provenir
des diflficultes de mensuration. Afin d'arriver a une certitude parfaite
nous avons mesure individuellement, en 1912, chaque plante et trace
des courbes dont I'examen ne laisse plus aucune incertitude sur la
repartition des plantes issues de naines en deux groupes bien distincts.

Je reproduis ci-dessous quelques-unes de ces courbes. (Fig. 1 a 3.)

Philippe de Vilmorin

73

Fig. 4. (Descendance d'une plante grande.)

II y a des cas cependant oii les deux groupes se confondent plus ou
moins et ou il devient difficile de dire qu'une plante appartient a la
cat^gorie des gi-andes ou a celle des naines. (Fig. 5 et 6.)

74

Snr une Race de BJe Nain Infixable

Fig. C.

Ceci est du a la fluctuation qui se manifeste dans tous les bl6s,
niais pas plus dans cette vari6t6 que dans les autres ; pour m'en assurer
j'ai fait des mensurations sur des varietes fix6es, mesurant des plantes
d'un meme lot, qui descendaient toutes d'un meme parent de 1910 et
avaient 6t6 cultivtes dans le meme terrain.

Voici, par exemple, la courbe trouvee pour le ble de Bordeaux
(Fig. 7).

180

160

150
Fig. 7.

120

Philippe de Vilmorin 75

De nieme, j'ai troiive que la taille, dans iin lot homogene de ble
blanc a paille raide, variait de 88 centimetres a 1 m. 50, dans le ble de
Noe de 1 m. 04 a 1 m. 54, dans le Victoria d'automne 1 m. 40 a 1 m. 94;
or, dans le ble que nous etudions, la difference moyenne entre les
{Dlantes grandes et les plantes naines n'est que de 30 centimetres'. II
n'y a done rien detonnant a ce que les conditions d'ambiance soient
suffisantes pour faire passer une plante dans une categorie a laquelle
ello n'appartiendrait genetiquement pas.

C'est evidemment ce qui s'est passe en 1904 pour le ble Shirno
(tamille blanche) ainsi qu'en 1905, ou une ou plusieurs plantes con-
siderees comme naines etaient genetiquement grandes, ce qui explique
les proportions tout a fait anormales de ces deux annees.

On ne peut etre absolument sur qu'une plante est reellement naine
qu'en etudiant sa progeniture. Toutefois il convient d'ajouter que
I'erreur possible provenant de ce chef est extremement faible et peut
etre pratiquement negligee.

Ce cas est done comparable a celui des souris jaunes. II est tres
possible que la conjonction de deux gametes portant le facteur nain
se produise reellement, mais ne puisse donner naissance a un germe.
Ceci expliquerait que les grains sont moins nombreux dans les epis
des plantes naines que dans ceux des plantes grandes ; mais si en
realite nous ne trouvons pas la proportion 2 a 1, c'est justement a
cause du non d6veloppement de certains grains, par suite duquel les
fleurs du centre de I'epillet peuvent developper leurs grains alors que
dans les conditions normales elles resteraient steriles.

Comme nous I'avons deja dit, s'il y avait reellement repulsion entre
les gametes portant le facteur nain, les ovules portant ce facteur
seraient fecondees par des gametes males depourvus du facteur, le
nombre de grains de pollen etant illimite. Dans ce cas nous aurions
une proportion de 3 plantes naines a 1 plante grande ; c'est I'hypothese
qui avait ete formulee au debut par Cuenot dans le cas des souris
jaunes, mais les recentes experiences de Miss Durham semblent bien
demon trer que la proportion est plutot 2 a 1 que 3 a 1.

Pour terminer, je signalerai que, si le caractere nain est dominant
sur le caractere grand dans cette famille, il n'en est pas de meme par

1 Toutefois dans quelques cas dans la descendance de plantes naines on trouve en
dehors d'un groupe de plantes grandes et d'un autre plus important de plantes naines
quelques individus de taille encore moindre variant de 30 a 60 centimetres. L'etude de
ces exceptions est rendue difficile par le fait que generalement ces plantes sont steriles.
Nous en avous cependant plusieurs en observation dont le produit sera examine I'annee
prochaine.

"6

Stir If lie Race de BU Nain Infixable-

rapport uu caractere grand des autrcs races de ble. C'est aiiisi qu'iin
croisement entre un ble Shirno nain, de 90 centimetres, et Ic ble Eclipse,
vari^te de 1 m. 50 de haut, a donne un F, homogene et compose de
plantes de 1 m. 20 a 1 m. 30 de haut.

Lc F., est represente par la courbe ci-dessous (Fig. 8).

yrv4

130

Fi". 8.

Dans un croisement entre une plante naine et une plante grande
du Shirno j'ai trouve en F„ sur ti plantes 1 grande et 5 naines '.

LITERATURE.

(1) Ca^NOT, L. Arch, de Zool. Exp. et Gen., Vols. I, n, in, vi.

(2) Durham, F. M. " Further experiments on the Inheritance of Coat Colour in

Mice" (Journal of Genetics, Vol. i, No. 2 (1911), p. 166).

1 Ces nombres sont malheureusement trop faibles pour avoir une valeur demon.strative.
Si rhypothfese envisagee est vraie on doit obtenir le meme nombre de nains et de grands.
Des croisements sur une plus grande echelle devrout etre tentes avant de couclure avec
certitude.

JOURNAL OF GENETICS, VOL. IN. NO. 1

PLATE II

CAMBRIDGE UNIVERSITY

PRESS r r <r

Bulletin

April MCMXIII

O. XXIX

Notes on new books and
journals published at the
Cambridge University Press,
Fetter Lane, London, E.C.
C. F. Clay : Manager

CONTENTS

OF

BULLETIN. No. XXIX

Theology

Twelve Parochial Sermons by J. E. B. Mayor

Twelve Cambridge Sermons by J. E. B. IVIayor

The Book of Judges (Cambridge Bible for Schools)

The Book of Ruth (Cambridge Bible for Schools) .

The First Book of Samuel (Revised Version for Schools) . . 26

Horae Semiticae, No. X

English

The Pronunciation of English in Scotland 9

French

A French Note-Book 26

Literature-

The Jataka, Inde.\ volume 10

Classical History

Greek Ohgarchies . 10

The Fine Arts

Byzantine and Romanesque Architecture II

Medieval Figure-Sculpture 12

History

The Cambridge History of India i

The Life of Philip Yorke, Earl of Hardvvicke 3

Early Wars of Wessex 3

British Borough Charters 4

The Colonization of Africa 26

Europe and the Far East ......... 26

The .Australasian Colonies ......... 28

Spain (1479— 1788) 26

Economics

The Economics of Everyday Life ....... 5

Law

The Panama Canal Conflict "... 13

Roman Private Law 14

Cambridge Manuals 24

Cambridge Naval anh Military Series 6

Continia-d on p- }, of wrapper

The Cambridge History of India. Under the editorship
of Professor E.J. Rapson, M.A., Lieut-Col. T. W. Haig,
C.M.G., and Sir Theodoi'e Morison, K.C.I.E.

In six volumes. Royal 8vo. Buckram, gilt top. With a large number of maps
and illustrations. Price 15J. net per volume; to subscribers I2.t. net.

The Syndics of the Cambridge University Press have
undertaken to publish a comprehensive History of India from
the eariiest times down to the present day. The work will
thus cover a period of more than three thousand years and
will deal concisely with the political and social history of the
numerous nationalities which have risen to power in the
Indian continent during historical times. Hitherto the three
main periods of Indian History — Ancient Hindu, Muham-
madan and British — have usually received separate treatment.
There is no single work in existence which deals adequately
with the subject of Indian History as a whole. The Syndics
trust that, by securing the cooperation of a number of scholars
under the direction of an editor for each of the main periods,
they will be able to supply the undoubted need for a standard
History of India.

As in The Cambridge Modern History, the aim of each
chapter will be to summarise in an accurate and readable
manner the results of the latest investigations in the subject
with which it deals ; and to each chapter will be appended a
select bibliography of authorities. The work will thus satisfy
the requirements both .of the general reader and of the
student.

The maps, however, will be inserted in the text and not
gathered together in a separate volume ; and the volumes
will be abundantly illustrated. As the monuments of India,
its inscriptions and coins, supply the most valuable historical
and chronological information, special chapters will be devoted
to these sources of history ; and photographs or other repro-
ductions of the most noteworthy objects of interest will be
added. For the means of supplying a larger number than
would otherwise have been possible of these and similar
illustrations, the Syndics are indebted to the generosity of
Sir Dorab Tata.

THE CAMBRIDGE HISTORY OF INDIA— Continued

The work will be complete in six volumes, royal 8vo, of
between 500 and 600 pages each. The following is a general
summary of the character and contents of the volumes : —

THE EARLY HINDU PERIOD,
under the editorship of Professor E. J. Rapson, M.A.
Vol. I. Ancient India, from the j Vol. II. Medieval iNOiA^from about
earliest historical times to about

the beginning of the Christian era.
This volume will deal with the
earliest history of India as known from
its literatures and monuments. It will
include an account of the Persian and
Greek dominions in N. Western India,
of the Maurya empire, and of the various
kin£;doms which arose on its decline.

the beginning of the Christian era
to the .Muhamniadan conquests in
the I Ith century a.d.
The chapters in this volume will deal
chiefly with the empires of the Kushanas
and the Guptas : the Hun invasions ; the
empire of Harshadeva ; and the inde-
pendent kmgdoms in Northern and
Southern India.

THE MUHAMMADAN PERIOD,
under the editorship of Lieut. -Col. T. W. Haig, C.M.G.

Vol. III. Turks and Afghans, from

the end of the loth century .■i.D. to

the conquest of Northern India by

Babar, A.D. 1525.

This volume will deal with the

conquests of MahmQd of Ghazni and

of Muhammad bin Sam of Ghor and his

lieutenants ; the Slave, KhiljT, Tughlaq,

and Sayyid dynasties of Delhi, and the

rise of mmor local dynasties during the

decline of the power of the Tughlaq

dynasty. It will include an account of

Muhammadan conquests in Bengal.

Vol. IV. The Mughal Empire,
from a.d. 1525 to 1757.
This volume will contain an account
of the establishment of the Mughal
empire by Babar, its temporary eclipse,
and its extension and consolidation by
Akbar, of the gradual e.xtinction of the
independent dynasties of the south by
Akbar, Jahanglr, Shahjahan, and Au-
rangv.ib, the rise of the Marathas and the
decline of the Mughal empire down to
the battle of Plassev.

BRITISH INDIA,
under the editorship of SIR THEODORE MORISON, K.C.I.E.

Vol. V. The Honourable East
India Company.
This volume will give an account of
the various European settlements in
India and of the history of the East
India Company from its foundation in
1600 a.d. to the Indian Mutiny in 1857.

Vol. VI. India under the Crown.
The distinctive feature of this volume
will be an attempt to estimate the im-
portance of those political, religious,
social and economic developments
which are principally due to British
rule.

It is hoped that the first volume will be published by
Easter 19 14, and that the remaining volumes will appear
in chronological order at regular intervals of six months.

Terms to subscribers
The subscription price for the whole work will be ^3. \2s.
net ; payable in six instalments of i 2^-. net each, one instalment
becoming due on publication of each of the volumes.

2

The Life and Correspondence of Philip Yorke,
Earl of Hard-wicke, Lord High Chancellor of
Great Britain. By Philip C. Yorke, M.A. Oxon.,
Licencid-es-Lettres of the University of Paris.

Royal 8vo. In three volumes. With 6 illustrations. Vol. I. pp. -\vi + 686.
Vol. II. pp. viii + 598. Vol. III. pp. viii+654. Price 45 j-. net.

Publishers Note

This book Is based on the Hardwicke and Newcastle
MSS and, in addition to the description of the life of Lord
Hardwicke, gives the whole history of the Georgian period
from 1720 to 1764. An account of the great judge's work in
the King's Bench and in Chancery is included. The characters
and careers of Walpole, Newcastle, Henry Pelham, the elder
Pitt, Henry Fox, the Duke of Cumberland, George H and
George HI, and various incidents — such as the fall of Walpole,
the Byng catastrophe, and the struggle between George 1 1 1
and the Whigs — appear in a clearer light, which, aid.ed by
original papers and MSS, the author has been enabled to
throw upon them. These documents are now published, or
brought together and annotated, for the first time.

Early Wars of Wessex. Studies from England s First
School of Anns in the West Country. By Albany F.
Major, Atithor of Sagas and Songs of the Norsemen,
etc., Edited by Charles W. Whistler, M.R.C.S., Atithor
of K Thane of Wessex ; King Alfred's Viking, etc.

Demy 8vo. With maps, plans and diagrams. Price 10s. 6d. net.

Publishers Note

Early Wars of Wessex is the title of a series of studies in
the history of the pre-Norman period. Their distinguishing
feature is that, while dealing only with the warfare that took
place in a comparatively limited area between Saxon and
Briton and Saxon and Dane, they show, nevertheless, that
these "battles of the kites and crows," as Milton con-
temptuously called them, had a far-reaching influence on the
future of England.

[A special prospectus of this book ivill be foT-warded on application^

3 1-2

British Borough Charters^ 1042 — 1216. Edited by
Adolphus Ballard, LL.B. {Loud.), Hon. M.A. (Oxon.),
Town Clerk of Woodstock, Author of The Domesday
Boroughs and The Domesday Inquest.

Royal 8vo. pp. cxlviii + 266. Price 15.?. net.

Extract from the Preface

In these pages I have collected about 330 charters and
documents relating to the boroughs of the British Isles, which
can be dated before the death of King John, and have analysed
them into their constituent clauses, and rearranged them in
the form of a code. The three parts of the Introduction will
furnish all necessary explanations ; the first part explains our
methods of study and specifies the sources from which the
documents are derived : the second part is a short commentary
on the code : the third part contains four supplemental essays,
in two of which I have attempted to find some general
principles underlying the code of municipal privileges, while
in a third I have compared some contemporary foreign
municipal charters with those of the British Isles. But in
these essays, I have dealt only with the problems of the
twelfth century, except where express reference is made to
other periods : and it does not follow that, because I have
hazarded certain opinions with regard to the formation or
characteristics of boroughs in that century, I hold the same
opinions with regard to the same problems either in earlier or
later centuries.

CONTENTS
Introduction.

I Preliminary. II Notes on the Code. Ill Supplemental Essays.

CHARTERS

I Formation of Borough.

IIa Burgage Tenure and Law of Real Property.

IIb Tenunal Privileges.

III Burgess Franchise.

IV Jurisdictional Privileges.

A. Courts. B. Modes of Trial. C. Procedure. I). Punish-
ments. E. Distress.

V Mercantile Privileges.

A. Markets and Fairs. B. Guilds and Trades.

VI Borough Finances.

VII Borough Officers.

.\|jpendix. (a) Additional Charters ; (/') Addenda et Corrigenda.
Index.

The Economics of Everyday Life. A First Book of
Economic Study. Part I. By T. H. Penson, M.A.,
Lecturer in Modern History and Economics at Pembroke
College, Oxford.

Large Crown 8vo. pp. xiv+176. Price y. net.

Extract from the Preface

Political Economy, or as it is more generally called at the
present day, Economics, is a subject which has hitherto found

very little place in a general education As a matter of

fact, however, the subject is an intensely practical one ; it
deals with problems of everyday life, and everyone can find
abundant material for observation and comparison in his own
home and in his own surroundings. Moreover it is being
increasingly recognised that for the proper discharge of the
duties of citizenship some knowledge of Economics is abso-
lutely essential. Great social and economic problems are
constantly needing solution, and those who are called Upon
to assist in that solution by supporting or opposing any par-
ticular measure or line of action should have a sufficient
understandine of economic cause and effect to be able to act
with judgment and conviction. The foundation of such
economic study may well be laid during the period of school
life, but for this a text-book is necessary in which the essential
features of the subject are simply described and clearly ex-
plained. An introductory book of this kind is besides often
felt to be a necessity even by those of more mature years,
social workers and others, who are taking up the subject for
the first time. The present work is an attempt to supply
both needs.

Part I contains two introductory chapters, and deals with
Production, Exchange and Distribution. Part II will deal
with Consumption, Taxation, and Trade Unions and Co-
operative Societies.

Morning Post. — It is becoming daily more clearly recognized that some
knowledge of economics is essential for every citizen, and that we need
books in which the treatment is not too difficult or too far removed from
the ordinary experience of the young student. The present small volume
is an admirable book of this kind. It is clear, concise, and practical in
its methods and illustrations, and it is strictly elementary in its character.
All schoolmasters ought at least to see and consider it.

The Cambridge Naval and Military Series. Under
the general editorship of Julian S. Corbett, LL.M.,
F.S.A., of Trinity College and the Middle Temple,
Lecturer in History to the Royal Naval War College;
and H. J. Edwards, C.B., M.A., Fellow and Tutor of
Peterhoti.se, and Colonel Commanding the Cambridge
University Contingent, Officers Training Coifs.

Demy 8vo. About 300 pages in each volume. With diagrams, maps,
plans, etc. The prices will vary from ^s. net to lo.f. net.

Having regard to the increasing demand in both Services
for standard works on naval and military subjects, and to the
tact that, in response to the growing recognition of their
importance and interest on all sides, such subjects have been
definitely included in the curriculum of the University, the
Syndics of the Cambridge University Press have decided to
publish a Naval and Military Series.

It will be their object, in issuing these works — of which
a first list is given below — to meet the needs of students both
in and out of the Services as well as those of the general
reader. The more technical books will be concerned with
strategy, tactics, geography, map-reading, armament, adminis-
tration, combined operations and the economics of war. The
volumes of a less technical, or non-technical, character will
deal with the wide range of subjects that appeal to all who
take a serious interest in Naval and Military affairs, including
studies of separate campaigns, biographies of commanders and
statesmen distinguished in the conduct of war, the history
and antiquities of the Navy and Army, and the political
aspects of Imperial Defence.

The following vohimes have already been arranged : —

A. Naval.
The Navy in the War of 1739—48. By Captain H. JF. Richmond, R.N.
Ocean Transport and Shipping. By Douglas Owen, Lecturer at the Royal

Naval War College.
Maritime International Law. By A. Pearce Higgins, M.A., LL.D.

B. Military.
Strategy for Englishmen. Bv Brevet Lieut-Colonel W. D. Bird, D.S.O.
From Bull Run to Port Republic, 1861—2. By Lieut.-Colonel W. K.

Seharlieh.
Non'Regular v. Regular Troops in the North of France, 1870 — /.

By Brevet Lieut.-Colonel L.J. Bols, D.S.O.

6

The Cambridge Public Health Series. Under the
general editorship of G. S. Graham- Smith, M.D.,

Lhiiversity Lecttirer in Hygiene, and Secretary to the
Sub-Syndicate for Tropical Medicine, of Pembroke
College; and J. E. Pzirvis, M.A., University Lecturer
in Chemistry and Physics in their application to Hygiene
and Preventive Medicine, and Secretary to the State
Medicine Syndicate, of St Johns and Corpus Christi
Colleges.

Demy 8vo. From 250 to 350 pages m each volume. The prices
will vary from -^s to \05. bd. net.

In view of the increasing importance of the study of
public hygiene and the recognition by doctors, teachers,
administrators and members of Public Health and Hygiene
Committees alike that the sains populi must rest, in pare at
least, upon a scientific basis, the Syndics of the Cambridge
University Press have decided to publish a series of volumes
dealing with the various subjects connected with Public
Health. Such a series has, indeed, a wide field, of which the
extent may be seen from the following provisional lists of
subjects, all of which will be treated by experts.

A. The Causation of Tuberculosis Serum Diagnoses

Biting Flies and Disease The Bacteriology of Foods

House Flies and Disease Methods of Post-Mortem Ex-

Ticks and Disease aminations

B. Sewage Disposal Tropical Hygiene

Water Purification Fever Hospital .Administration

School Hygiene Sanitary Law and Practice

Physical Education Sound and Unsound Foods

Ventilation Domestic Sanitation

Soils, Subsoils and CHmate in Offensive and Noxious Trades

relation to Health Chemical ."Analyses of Foods

Thus the whole series will appeal not only to medical men
but also to those engaged in the study or administration of
public health at home or abroad, and to those who are in-
terested in Public Health matters, as e.g. County and Town
Councillors. The volumes in the first of the above sections
will be useful to laboratory students, colonial doctors and
bacteriologists in particular ; those in Section B to civil and
municipal engineers, architects, school-teachers, medical officers,
public analysts, and sanitary officers.

Cambridge Bible for Schools and Colleges. General
Editor for the Old Testament : A. F. Kirkpatrick, D.D..
Dean of Ely.

Extra fcap. 8vo. With introductions and notes.

The Book of fudges. In the Revised Version. Edited by
G. A. Cooke, D.D., Hon.D.D. Edin., Oriel Prof ssor of
the Interpretation of Holy Scripture, Oxfrd, and Canon
of Rochester, Canon of St Mary s Cathedral, Edinburgh.
pp. xlii-\- 204. With a map. Price is. net.

The Book of Ruth. In the Revised Version. Edited by
G. A. Cooke, D.D. pp. xviii+ 22. Price is. net.

The Books of fudges and Ruth. In the Revised Version.
As aboz'c, in one volume. Price 2s. 6d. net.

Txvelve Parochial Sermons. By fohn E. B. Mayor,
M.A., F.B.A., late President of St fohn s College and
Professor of Latin in the University of Cambridge.

Crown 8vo. pp. viii + io8. Price is. bd. net.

Extract fro7n the Preface

This volume is issued in fulfilment of a promise made
twelve months ago in the preface to John Mayor's Twelve
Cambridge Sermons. It will exhibit him in a fresh light to

those who knew him only as scholar and professor It has

sometimes been supposed that Mayor when he preached

could not bring himself down to the level of his hearers

But that he was incapable of using his knowledge in a

way that commended itself even to the unlettered is an entire
mistake, as these sermons prove.

Church Times. — Whether we regard the thoughts, or the diction, or the
method of these sermons, there is one word that exactly expresses their

character — they are beautiful No one can read the third sermon, on

"The Desire of All Nations," without feeling that he has come into a
new atmosphere, invigorating, fresh, and inspiring. All preachers,
whether learned or unlearned, can take a lesson from Prof. Mayor's style
in the art of presenting profound truths in a homely but always reverent
way.

Tivelve Cambridge Sermons. By fohn E. B. Mayor,
M.A., F.B.A. Edited, with a memoir, by H. F. Stewart,
B.D., Fellow and Dean of St fohn s College.

Crown 8vo. pp. lxviii + 254. Witli photogravure portrait. Price 5^. net.

The Commentaries of Isho^dad of Merv^ Bishop
oj Hadatha (c. 850 A.D.). Vol. IV, Acts oj the
Apostles and Three Catholic Epistles in Syriac
and English. Edited and translated by Margaret
Dimlop Gibson, LL.D. {St Andreius), Litt.D. {Dublin).
With an introduction by J. R. Harris, M.A.

Horae Semiticae, No. X. Crown 410. pp xvi + 96. With facsimile.
Price IS. i>d. net

Extract from the Editor s Preface

I have adopted for the text of this volume that of the MS
kindly lent me by the Rev. Professor D. Margoliouth of
Oxford, called in previous volumes Codex M, as several
critics have wisely pointed out its very decided superiority
to Dr Rendel Harris's MS (Codex H) which I have accord-
ingly relegated to the notes.

The Pronunciation of English in Scotland. By

William Grant, M.A., Lecturer on Plionetics to the
Provincial Committee for the Training of Teachers,
Aberdeen, Convener of the Scottish Dialects Committee.

Cambridge Primers of Pronunciation. Crown 8vo. pp. xvi + 208.
Price y. 6d. net.

Extract from the Preface

This book is intended primarily as a Phonetic Manual for
the use of students in Scottish Training Colleges and Junior
Student Centres, but it is hoped that it may prove useful to
teachers of English of all grades in our Scottish schools, and

to all who have to engage in public speaking Part I deals

with the manner and place of iormation of the various sounds

and their chansjes in combination It also enumerates the

variations from Standard speech and gives suggestions for
the correction of errors of pronunciation. Part II consists of
a series of texts written in the speech of the educated middle
classes of Scotland. Part III contains a series of questions
on the subject-matter of Part I. The Appendix contains
(1) the ordinary English spelling of the phonetic texts in
Part II, (2) an account of the chief differences between
Scottish and Southern English, (3) advice to teachers on the
subject of the teaching of reading.

9 i-S

The Jdtaka, or Stories of the Buddha's Former
Births, TrcDislated from the Pali by Various Hands.
Index Volume.

Royal 8vo. Buckram, pp. iv + 64. Price 5.S. net.

The publication of the inde.x volume completes this work. The stories are

contained in six volumes, price I2.r. dd. net each; the price for the set

including the index volume, Ijeing ^3. y. od. net.

Extract from the Preface to J^ohtme I

It is quite uncertain when these various birth-stories were
put together in a systematic form such as we find in our

present Jataka collection Some of the birth-stories are

evidently Buddhistic and entirely depend for their point on
some custom or idea peculiar to Buddhism ; but many are
pieces of folk-lore which have floated about the world for
ages as the stray waifs of literature and are liable everywhere
to be appropriated by any casual claimant.

The Jatakas themselves are of course interesting as speci-
mens of Buddhist literature ; but their foremost interest to us
consists in their relation to folk-lore and the light which they
often throw on those popular stories which illustrate so vividly
the ideas and superstitions of the early times of civilisation.
In this respect they possess a special value, as, although much
of their matter is peculiar to Buddhism, they contain em-
bedded with it an unrivalled collection of Folk-lore. They
are also full of interest as giving a vivid picture of the social

life and customs of ancient India They form in fact an

ever-shifting panorama of the village life in the old days
before the Muhammadan conquest.

Greek Oligarchies. Their Character and Organisation.
By Leonard IVhibley, M.A., Felloiv of Pembroke College,
Cambridge.

Reissue. Crown 8vo. pp. viii-f2l2. Price 3^-. td. net.
CONTENTS

CHAP.

I The Classification of Constitutions : The Claims and Character of

Oligarchy.

II The Causes of Constitutional Change.

III The Historical Development of Constitutions.

IV Varieties of Oligarchy.

V Organisation of Oligarchic Government.
Index.

10

Byzantine and Romanesque Architecture. By

Sir Thomas Graham Jackson, Bart., R.A., Hon. D.C.L.
Oxford, Hon. LL.D. Cambridge, Hon. Fellow oj
Wadham College, Oxford.

Crown 4to. In two volumes. Vol. I, pp. xx + 274. Vol. II. pp. viii + 286. With

165 plates, 4 of which are coloured, and 148 illustrations in the text, a large

number being reproduced from the author's own drawings. Bound in cloth, with

parchment back lettered and ornamented in gold, gilt top.

Price £2. zs. od. net.

EXTRACTS FROM PRESS NOTICES

A/kenaeum. — What an accomplished architect writes on the history of his art
must always be interesting, and Sir Thomas Jackson long ago proved

that he is a scholar as well as an artist The author's aim is to instruct

by the study of the past, to explain as well as to describe. All that, as
one turns over the two handsome volumes, is obvious ; it is obvious also

that the aim is achieved He has produced a very interesting as well

as a beautiful book.

Outlook. — -In two handsome volumes, profusely illustrated by drawings,
photographs, and diagrams. Sir Thos. Jackson has traced the history of

Byzantine and Romanesque styles to the advent of the Gothic The

book has literary qualities of a very high order, and is one of the most
readable we have ever read. A master of his subject, Sir Thos. Jackson

has a gift of clear and felicitous expression This fine work cannot be

praised too highly.

Britisti Arcliited. — By his simple, lucid style and the wide range of his

subjects Sir T. G. Jackson has provided a large and most valuable

contribution to the history of architecture, which we can hardly imagine
is likely to be displaced by anything else. The whole format of the
volumes is excellent, the size and shape of them, the beautiful type, and
the excellent illustrations.

Westminster Gazette. — Sir Thomas Jackson's drawings are a charming com-
promise between architect's and artist's work So with his comment.

He gives us what is in effect a caialogjie raisojinl oi the chief buildings —
a thing invaluable to the student — but he is mercifully sparing of techni-
calities, and the whole work may be read with pleasure.

Morning Post. — Sir Thomas Jackson always writes not only like a practical

architect but like a scholar and a connoisseur His wide knowledge of

history and literature enables him to set the subject he is dealing with in
its context, as it were, and to view it as part of the scheme of culture of

that epoch These two volumes are worthy of the close attention of

everyone interested in architecture.

Standard. — Sir Thomas Jackson writes with not less enlightenment than

authority To the general reader wdth only the remotest knowledge

of architecture the book cannot fail to be absorbingly interesting.

\^A special 8 //. prospectus, with specimen pages and plates, will
be forwarded on application^

I I

An Account of Medieval Figure- Sculpture in
England. By Edward S. Prior, M.A., F.S.A.,
Slade Professor of Fine Art in the University of
Cambridge, and Arthur Gardner, M.A., F.S.A.

Uemy 4to. pp. xii + 734. With 855 photographs. Price ^3. y. od. net.

EXTRACTS FROM PRESS NOTICES

Athenaeum. — Prof. Prior and Mr Gardner have given us a work on medieval
figure-sculpture which will surely mark an epoch in the history of English
art. On the lowest ground it is an enchanting picture-book, with its
855 photographs — many of them of masterpieces of sculpture — illus-
trating the often unnoticed wealth of Gothic art in this country

This is the work of men with a thorough knowledge and love of their

subject It is incumbent on every lover of English Gothic art to

familiarize' himself with it.

Studio. — The joint authors of a book that, even without its deeply interesting
text, must be a delight to all lovers of the noble art of decorative figure-
sculpture, on account of the vast number and the beauty of its illus-
trations, go to the very root of the matter under discussion. Not only
do they describe and classify all the most characteristic examples of this

delightful craft that still survive in England they realise the very

spirit that animated those who executed them.

Times. — Taken as a whole the book deserves that prodigally misused epithet

of epoch-making If it were nothing more, and it is a great deal more,

" Medieval Figure-Sculpture in England " would be an unrivalled picture-
book. Merely turning over the pages at ha<?ard, the reader lights on

fresh and delightful things Neither amateur nor student could wish

for better guidance than that which Professor Prior and Mr Gardner have
provided.

British Architect. — The student of Gothic architecture may now pass before
him a comprehensive review, a splendid panorama of examples of the
best periods of medieval sculpture in this country by means of the fine

volume on "Medieval Figure-Sculpture in England." Our medieval

figure-sculpture may now, for the first tim.e, be appreciated at something
of its real value.

Architect. — A sense of the amplitude of medieval art is the dominant impres-
sion that we gather from a first perusal of the masterly work on Enghsh
figure-sculpture in the middle ages with which Mr Prior and Mr Gardner

have endowed us The authors have produced a splendidly complete

conspectus of medieval figure-sculpture in England in all its branches.

Guardian. — It would be difficult to overpraise this work Not a point

that we can detect is passed over. Every stage of English medieval
sculpture is noticed, discussed, classified, explained. A wealth of illus-
trations is provided, carefully arranged so as to adjoin and elucidate the
text to which they relate.

\A special 8 pp. prospectus with specimen plates will be sent on application^

12

The Panama Canal Conflict betxveen Great Britain
and the United States of America. A Study by
L. Oppenheim, M.A., LL.D., Whewell Professor of
Internatio7ial Laiv in the University of Cambridge,
Member of the Institute of Internationa/ Law.

Crown 8vo. Second edition, pp. iv + 5S. Price 2s. bd. net.

Exti^act prom the Preface

This study had been written before the diplomatic corre-
spondence in the matter was available and without further

information than that which could be gathered from the
Clayton-Bulwer Treaty, the Hay-Pauncefote Treaty, the
Hay-Varilla Treaty, the Panama Canal Act, and the Memo-
randum which President Taft left when signing that Act.
Hence, the reader is presented with a study which is abso-
lutely independent of the diplomatic correspondence, and he
can exercise his own judgement in comparing my arguments
with those set forth /ro et contra the British interpretation of
the Hay-Pauncefote Treaty in the despatches of Sir Edward
Grey and Mr Knox.

Times. — It contains a lucid statement of the British case ; of the bearing of
the controversy on the practice of the United States respecting the
relations between International Law and their Municipal Law ; and of
the position with regard to arbitration and the British American Arbi-
tration Treaty.

Scotsma?t. — It is a thoughtful and instructive study of as interesting a case as
ever came up in the long history of international law, and will prove
profitable reading to students on both sides of the Atlantic who wish to
reach well-founded views of the interpretation of the clause in the Hay-
Pauncefote Treaty that has given rise to the dispute.

Economist. — The Whewell Professor of International Law at Cambridge
discusses the legal aspects of the Panama Canal controversy, and argues
against the contention of President Taft that the words "all nations"

mean all foreign nations This well-written little book deserves to

be widely read.

Cambridge Review. — Professor Oppenheim's authoritative pronouncement on
the subject is of much interest and importance. The exemption of
vessels engaged in the coasting-trade of the LTnited States from the pay-
ment of tolls raises two vital questions. How far will it damage our
shipping-trade ? Does it involve a breach of the Hay-Pauncefote treaty ?
Prof. Oppenheim does not concern himself with the first question,

which is economic rather than legal But the breach of a treaty is in

itself a serious matter, and the second question must therefore be faced
whatever answer is given to the first. Prof. Oppenheim has dealt with
it in a lucid and, in our opinion, conclusive manner.

13 1—7

Elementary Principles of the Roman Private Laiv.

By W. W. Buckland, M.A., Fellozv and Tutor of Gon-
ville and Cuius College, Cambridge.

Demy 8vo. pp. viii + 420. Price \os. bd. net.

EXTRACTS FROM PRESS NOTICES

Juridical Review. — The pre.sent volume is a handbook for students of the

Itistitutes of Gaius and Justinian. In achieving this object it follows a
somewhat novel but very useful method, being neither a textual com-
mentary on those ancient manuals nor a complete exposition of their
contents. The author assumes a knovvledge of the text, and discusses
the principles involved in the doctrines there laid down, presenting in a
luminous and suggestive manner the difficulties which arise and the

current theories as to their solution The book is sane and sound

throughout Mr Buckland tries to present the law from the point of view

of the practising Roman lawyer. The result cannot fail to stimulate
the interest of the student and help him to realize that the law of Rome
was not a collection of dry bones, but a living, full-blooded reality.

Lmv Magazine. — This is a scholarly and thoughtful treatise on the inex-
haustible subject of Roman law The practitioner who wishes to

revive his learning, and the student who is preparing for examination
will find in this a valuable and ready help.

Law Quarterly Revieiv. — Mr Buckland has well fulfilled his purpose of
showing the Roman law as ' not merely a set of rules on paper, a
literary product, but a group of institutions under which the Romans
actually lived.' The spirit is as far as possible from that of the old-fashioned
manuals which combined exaggerated letter-worship of the Roman jurists

with misunderstanding, often gross, of the subject Altogether this is

a most commendable piece of work, and highly creditable to the Cam-
bridge law school.

Latv Times. — The book is specially designed for students who have read
their Institutes and but little more, and .VIr Buckland is to be con-
gratulated upon producing an exceedingly useful and interesting work.

Scotsman. — A student who is expected to know no more of Roman law than
Gaius and Justinian can teach him, might search long and vainly before
coming on a book better suited to his purposes. The commentary
supplements without attempting to supersede the old Institutes themselves,
and carries lightly a great weight of learning both in the old texts them-
selves and in the endless modern literature of their latter-day expositors.

It cannot but prove profitable reading to all sorts of students who

have for the first time to take up the Civil Law.

Oxford Magazine. — This is a learned and valuable book, which does what no

other work in English, so far as we know, attempts to do It does

not profess to be a systematic exposition of the Roman Law (though
there is admirable exposition in it), but a series of critical discussions of
Roman institutions, and the theories which have clustered round them.

Mr Buckland states that his object is "to suggest and stimulate

rather than to inform." This end he has achieved.

[4

Scientific Papers. By /. Y. Buchanan, M.A., F.J^.S.,
Comniandeur de /' Ordre de Saint Charles de Monaco,
Chemist and Physicist of the Challenger Expedition,
Vice-President dii Coniitd de Perfectionnenient de I'ln-
stitut Oc^anographicjne [Fondation Albert i"' Prince de
Monaco). Volinne I.

Demy 8vo. pp. xii + 314. With a portrait, maps, charts and diagrams.
Price los. 6d. net.

CONTENTS

I On the Absorption of Carbonic Acid by Saline Solutions. (1874.)

II Some Observations on Sea-water Ice. (1874.)

III Note on the Vertical Distribution of Temperature in the Ocean.

(1875-)

IV On the Determination, at Sea, of the Specific Gravity of Sea-water.

(1875-)
V Preliminary Report to Prof. Wyville Thomson, F.R.S., Duector of
the Civilian Scientific Staff, on Chemical and Geological Work
done on Board H.M.S. "Challenger." (1876.)
VI Distribution of Salt in the Ocean, as Indicated by the Specific Gravity

of its Waters. (1877.)
VII Preliminary Note on the Use of the Piezometer in Deep-sea Sounding.

(1876.)
VIII Laboratory E.xperiences on Board the " Challenger." (1878.)
IX and X Notes on the Distribution of Temperature under the Ice in

Linlithgow Loch. (1878-79.)
XI Deep-sea Investigation, and the Apparatus employed in it. (1881.)
XII and XIII On Oceanic Shoals and on Tidal Currents in the Ocean,

discovered in the s.s. " Dacia " in October, 1883.
XIV and XV On Land Slopes separating Continents and Ocean Basins and
on the Exploration of the Gulf of Guinea. (1887.)

Morning Fast. — The very valuable contributions to ordered knowledge pub-
lished from time to time by Mr J. Y. Buchanan, whose oceanographical
work has had such wide and far-reaching results, are now to be collected
and published. The first volume contains papers dealing with a part of
the work which Mr Buchanan did as Chemist and Physicist to the
Challenger Expedition, as well as with some later work, notably that
done in the Buccaneer.

Aberdeen Daily Journal. — As is well known Mr Buchanan's work is of a
fundamental character, and the publication of these memoirs m collected
form will serve a most useful purpose and is sure to be welcomed by all

investigators and others interested in the science of the sea An

excellent photograph of the author is given as a frontispiece.

A second volume is in preparation.

\A prospectus of this book will be sent on application]

15

Reports of the Committee on Electrical Standards
appointed by the British Association for the
Advancement of Science. A Record of the History
of "' Absolute Units" and of Lord Kelvin's work in
connexion zuitk these. Reprinted by permission of the
Council.

Demy 8vo. pp. xxiv + 784. With 10 plates and 46 text figures.
Price I2s. bd. net.

Extract from the Introduction

The original British Association Committee on Standards
of Electrical Resistance was appointed at the suggestion of
Professor William Thomson (later Lord Kelvin) in 1861
and consisted of Professor A. Williamson, F.R.S., Professor
C. Wheatstone, F.R.S., Professor VV. Thomson, F.R.S.,
Professor W. H. Miller, F.R.S., Dr A. Matthiessen, F.R.S.,
and Mr Fleeming Jenkin. The principal object of the
Committee was, first, to determine what would be the most
convenient unit of resistance, and second, what would be the
best form and material for the standard representing that

unit From 1862 to 1870 much valuable work was done

by the Committee which in the interim had been strengthened
by the addition of Sir Charles Bright, Professor J. Clerk
Maxwell, Mr C. W. Siemens, Mr Balfour Stewart, Mr C. F.
Varley, Professor G. Carey Foster, Mr Latimer Clark, Mr D.
Forbes, Mr Charles Hockin, Dr Joule, and Dr Esselbach.
In addition to dealing with the primary electrical standards
the Committee have also considered the subjects of platinum-
thermometry, thermal and magnetic units, and physical con-
stants in general. During the latter years of the Committee s
existence it promoted international uniformity in standards,
and for this purpose many experiments were undertaken at
the National Physical Laboratory. The appointment by
the London Conference of 1908 of an International Scientific
Committee of fifteen to direct work in connection with the
maintenance of electrical standards relieved the Committee of
much of its responsibility. The main objects for which it
had been appointed had been achieved ; in all the principal
countries of the world the same units of resistance, of current,
and of electromotive force had been adopted and the standards
in use were practically identical.

16

Matrices and Determinoids. Volume I. By C. E.

Cnllis, M.A., Pi'ofessor of Mathejnatics in the Presidency
College. Calctdta.

b ■

University of Calcutta Readership Lectures. Volume I. Large Royal 8vo.
pp. .\ii+43o. Price z\s. net.

Extract Jrom the Preface

The chief feature of the present work is that it deals with
recta^icrular matrices and determinoids as disting-uished from
square 7natrices and detei'minants. An attempt is made
to set forth a complete and consistent theory or calculus of
rectangular matrices and determinoids.

The first volume contains the most fundamental portions
ot the theory, and concludes with the solution of any system
of linear algebraic equations, which is treated as a special
case ot the solution ot a matrix equation of the first degree.

A second volume, which is nearly ready, will contain
further developments of the general theory, including a dis-
cussion of matrix equations of the second degree. It will
also contain a large number of applications to Algebra and
to the Analytical Geometry of space of two, three and n
dimensions.

Principia Mathematica. Volume III. By A. N.

Whitehead, Sc.D., F.R.S., and Bertrand Russell, M.A.,
F.R.S.

Large Royal 8vo. pp. x + 492. Price 2\s. net.

Extract from, the Preface

The present volume continues the theory of series begun
in Volume II, and then proceeds to the theory of measure-
ment. Geometry we have found it necessary to reserve for
a separate final volume. In the theory of well-ordered series
and compact series, we have followed Cantor closely, except
in dealing with Zermelo's theorem, and in cases where
Cantor's work tacitly assumes the multiplicative axiom.
Thus what novelty there is, is in the main negative.
Part VI, on the theory of ratio and measurement, on the
other hand, is new, though it is a development of the method
initiated in Euclid Book V and continued by Burali-Forti.

Mendel's Principles of Heredity. By W. Bateson,
M.A., F.R.S., V.M.H., Honorary Fellow of St Johns
College, Director of the John Junes Horticultural Insti-
tution.

Third impression. Demy 8vo. pp. xvi + 414. With 3 portraits,
6 coloured plates, and 38 figures. Price \2s. net.

Note to the third impression

In the past three years the progress of Mendehan analysis
has been very rapid, and certain chapters of this book,
especially those dealing with Coupling and Repulsion, and
with the Heredity of Sex, are in essential respects out of date.
Knowledge of these subjects is at present in a transitional
stage, and I have endeavoured in a series of brief appendixes
to acquaint the reader with the nature of the principal advances
made, while awaiting an opportunity of rewriting the book.

EXTRACTS FROM PRESS NOTICES

Gardeners Chronicle. — A new impression cannot fail to be welcomed

MendePs Principles of Heredity is already a classic. It marks a position of
stability towards which previous work is now seen to have logically con-
verged, and from which new and active research is to-day no less logically
diverging. The various waves of biological thought are constantly inter-
secting, mingling, and passing on with altered rhythm, but it rarely
happens that so many meet together at a nodal point as during the last

decade As an analysis of that point, as a picture of how it has come

into being, and as a foreshadowing of happenings in the near future,
MendePs Principles stands alone, and it is good to know that the
generation of students now growing up cannot be cut off from the
possession of a book so full of inspiration.

Lancet. — Professor Bateson's admirable book puts out in the clearest possible

manner the whole story of Mendeiism We have read this book with

the greatest possilile interest and recomnjend it to all our readers.

Journal of Botany. — The present work is the most complete treatise on the

Mendelian aspect of Heredity which has yet appeared in English

It is a privilege to have read Mr Bateson's work, and to have assimilated
the exposition of the principles which he so ably advocates, illustrated
with concise tables, as well as figures and coloured plates, which enhance
its value as a solid contribution to English Science.

Contemporary Review. — Mr Bateson's long-expected volume on Mendeiism
undoubtedly marks a new stage, probably a new era, in the investigation

of the origin of species It is, of course, impossible in so short a

notice to do more than draw attention to a volume that works out in
close detail applications to natural phenomena of Mendel's wonderful
discovery. We can, therefore, merely commend in general terms a work
nf the first order in the thought of our generation.

18

The Genus Iris. By William Rickatson Dykes. With
forty-seven coloured drawings by- F. H. Round, one
coloured plate of seeds by Miss R. M. Cardew a7id thirty
line draivings by C. W. Johnson.

Demy Folio, pp. viii + 246. With 48 coloured plates and 30 line drawings.

Bound in Ro.xburgh — dark green leather back and green cloth sides — with

lettering in gold and gilt top. Price ^6. 6^. od. net.

EXTRACTS FROM PRESS NOTICES

Times. — Mr Dykes has succeeded Sir Michael Foster as the chief authority
on the iris ; and, like Foster, he has not been content with a merely
botanical knowledge of it Besides examining the herbarium collec-
tions at Kew, Berlin, Vienna and other places, he has grown in his own
garden and, whenever possible, has raised from seed more species,
probably, than any one else in England, and perhaps than any one else

in the world This book must be the basis of all future study of the

subject for it is the result of knowledge both practical and theoretical,
and in that combination unique. The coloured drawings by Mr Round
are far more artistic in their simple accuracy than most modern drawings
of the same kind. The work does credit to the publishers as well as to

the author and artist [and] is remarkable among other works of the

same kind for its very precise and valuable cultural directions. Mr Dykes
gives precise directions for the raising of Irises from seed, and his chapter
on the suiiject is one of the most valuable parts of a valuable book.

Westminster Gazette. — It may be said at once that they [the publishers] have
amply merited the thanks of every horticulturist and botanist by the very
handsome style in which the vi'ork has been reproduced as regards text,

as well as illustration Mr Round, the illustrator of the book, has in

the treatment of the flowers come up to the task, and the colour-process
employed has rendered his beautifully delicate work with photographic

faithfulness The text is literally packed with information for the

botanist and for the grower of irises. Neither will be able to do without it.

Athenaeum. — Tb.is monumental work has a special interest, for it is not

exactly a botanist's monograph but rather a gardener's classification,

made, however, with strict regard to botanical usage Mr Round's

coloured drawings form a conspicuous feature of this magnificent volume.

We congratulate the Camliridge Press on the publication of this

important contribution to garden botany, and on the general production
of the volume.

Gardeners' Chronicle. — The author has for years been a keen collector and
successful grower of Irises. In this he has taken up and worthily
continued the late Sir Michael Foster's work. The result of it is

impressed almost on every page of the book Forty-eight folio

colour-plates, for the most part of exquisite execution in draughts-
manship and print, are no mean addition to a monograph of a single

genus The drawing and colouring of the flowers could hardly be

surpassed in delicacy, artistic finish, and faithfulness.

\A special prospectus of this hook, together with a specimen
coloured plate, ivill be forivarded on applicatio>i\

19

Herbals : Their Origin and Evolution. A Chapter
m the History of Botany, 1470- 1670. By Agnes Arber
{Mrs E. A. Newell Arber), D.Sc, F.L.S., Fellow of
Newnhani College, Cambridge, and of University College,
London.

Royal 8vo. pp. xviii + 254. With frontispiece, 21 plates, and 113 text-figures.

Price \os. hd. net.

EXTRACTS FROM PRESS NOTICES

Spectator. — This delightful book is a credit to Newnham scholarship and to
the Cambridge University Press. There never was a book richer in
pleasant old-world portraits and wood-cuts ; and there will never need to
be another book on herbals. Mrs Arber has made her book thoroughly
beautiful; the very initial letters of the chapters date from 1550. Among
the wood-cuts of herbs and flowers are many which show the utmost skill

of design Indeed, the beauty of the book endangers the text: and

reviewers who have souls above type will be so charmed with the
illustrations that they will forget to read what the book says. By which
forgetfulness they will miss a great wealth of scholarly, delicate, intimate
writing.

Manihester Guardian. — Mrs Arber has done a most interesting piece of work,

and done it with scholarly devotion and thoroughness Her work

is full of charming human touches. It could hardly have been better
done, and not the least admirable feature of the work is the illustrations
to it. The choice of these exhibits not only the judgment of a scholar

but the discernment of an artist also Mrs Arber's volume would be

worth possessing for the illustrations alone.

Scotsman. — A thoroughly interesting addition to the " books about books " is

made in Herbals: Their Origin and Evolution .Apart from the

attraction which so well executed a work has for those mterested in the
beginnings of botany, the illustrations make it a volume which every
lover of books, and especially of old books, will desire to possess.

Church Times. — This is a book which will appeal very much to all who

are interested in botany and its history and its connections with medicine.
It is a learned, exhaustive, and clear description of the herbals printed
during the two hundred years which followed the invention of printing

and of woodblock illustration No book of the kind has preceded

this, nor does it seem that room has been left for another of its kind.

Cambridge Magazine. — Mrs Arber has produced a volume written in a lucid
and attractive style which is not only a contribution of real importance
to the history of Botany, but a book that cannot fail to appeal to the

layman who has any artistic sense Mrs Arber's scholarly and

sympathetic treatment of the work of the Herbalists and of the artists
who collaborated with them is a fitting memorial to men who regarded it
a privilege to have been able with the help of God to make known to
their fellows the gospel contained in the book of Nature.

\A special prospectus oj this book, including a specimen plate, mav
be obtained on application']

20

Makers of British Botany. A Collection of Biographies
by living botanists. Edited by F. W. Olivej^

Demy 8vo. pp. viii + 332. With frontispiece, 26 plates and a text-figure.

Price gj'. net.

EXTR.ACT.S FROM PRESS NOTICES

Athenaeum. — This volume i.s the outcome of a course of ten lectures delivered
at University College, London, in the spring of 191 1, in which a series
of different botanists each gave the biography of one of the " fathers " of
the subject. To these biogra|)hies six chapters have been added, in

order to render the book "more fully representative." Each author

brings out well the value of the work of the old master he is representing,
and indicates its bearing on the science of the day. As a supplement —
and to some extent a corrective — to Sachs's classical "History of Botany,"
the present work must stand on the shelf of every serious botanist.

Gardeners' Chronide. — Professor Oliver has carried out a difficult and onerous
task with conspicuous success It is not often possible to give whole-
hearted praise to a book ; but such praise may be given to this botanical
anthology. We hope that it will be made a text book in our universities,
and that thereby our youth engaged in the study of botanical science
may be brought up in the knowledge and veneration of the men who
laid the foundations on which modern botanists are building.

Ftetd. — This volume consists of a series of sketches of the lives of eminent
British botanists treated in relation to their botanical labours and the
influence they have exerted on the development of the science of

botany Apart from the interest attaching to some of the biographies

on account of the strenuous lives and strong personalities they portray,
the historical review of the growth of botanical knowledge also contains

much that is both generally instructive and scientifically useful , the

book as a whole being eminently readable.

Aberdeen Journal. — No important aspect of the development of botanical

science is omitted The survey is both instructive and stimulating

This is a book which experienced botanists will find not only of genuine
interest, but full of suggestiveness with regard to the development of the
science, as all good histories are. Further, it is a book to be highly
commended to the attention of young botanists, who will not only find
the history of their subject pleasantly told, but who will be made to feel
the personal spell of those workers most of whose names they are alreadv
familiar with, and inspired to enthusiastic effort in their own held.

Glasgow Herald. — The book is valuable in bringing to notice the work of
some men who have really initiated sub-sciences and yet are not well
known. The articles are all by men specially qualified to appreciate and

expound their particular subject The book is well illustrated with

portraits of the botanists and reproductions of their drawings. There is
a comprehensive inde.x.

\_An illustrated prospectus of this book may be obtained on application'^

21

Vegetation of the Peak District. By C. E. Moss, B.A.
(Cantab.), D.Sc. {Vict.), F.R.G.S., F.L.S., Curator of
the Herbarium, University of Cambridge.

Demy 8vo. pp. x + 236. With 36 illustrations and 2 coloured maps in pocket.

Price 12^. net.

Extract from the Preface
The present work is the result of a botanical survey of the
Peak District of the southern Pennines begun in January,
1903 With regard to the nomenclature of plant com-
munities, the terms plant formation and plant association are
used in accordance with resolutions passed unanimously by
the British Vegetation Committee, and presented to the
International Congress of Botanists held at Brussels in

1 9 10 The names of plants are, as a rule, the same as

those given in the tenth, the latest edition of The London
Catalogue of British Plants (London, igo8). This being so,
the author-citation is omitted, as being unnecessary in a work
of this character ; synonyms, however, are added in special
cases. The sequence adopted is that of Engler's system,
which, in several European countries and in the United
States of America, is rapidly superseding that of Bentham
and Hooker.

CAMBRIDGE COUNTY GEOGRAPHIES

General Editor: F. H. H. Guillemard, M.A., M.D.

A series of County Geographies, price \s. 6d. each,
suitable for general use as hand-books to the various counties
and also intended for use in schools. Each volume gives an
account of the history, antiquities, architecture, natural history,
industries, and physical, geological, and general characteristics
of the county, and each has two coloured maps (one physical
and the other geological) and a large number of illustrations.

The latest addition to the series is : —

Herefordshire. By A. G. Bradley.

A list of the forty-six volumes already published, together
with prospectuses of particular volumes, will be sent on
application. Volumes on the remaining counties of England,
.Scotland, and Wales are in an active state of preparation ;
arrangements for a series of Irish geographies have also been
made.

22

The Duab of Turkestan. A Physiographic Sketch and
Account of Some Travels. By W. Rickmer Rickmers.

Large Royal 8vo. pp. xvi + 564. With 37 maps and diagrams, and
170 illustrations. Price 30.f. net.

EXTRACTS FROM PRESS NOTICES

Times. — Mr Rickmers is a geographical explorer of the highest qualifications.

There is very little ordinarily included within the province of

physical science which is not dealt with expertly in this book, and
nothing concerning mountains fails to be treated with the full knowledge

and sure touch of a master To an author who offers instruction,

amusement, vivid pen-pictures of unfamiliar and fascinating scenes and
such magnificent sun-pictures as the telescopic panorama of the glaciated
range of Peter the Great, one ought only to be grateful.

Daily Chronicle. — Mr Rickmers applies the name of Duab of Turkestan to
the land between the two rivers (du, two ; ab, water ; analogy, Punjab),
between the Amu-darya and the Sir-darya, or Oxus and Jaxartes. The
area contains everything that is typical of and common to various sub-
divided conceptions, such as Turkestan, Central, Middle or Inner Asia,

Turan, Iran, Bokhara, Kashgaria, Tarim, etc., etc Full of interesting

information of climate, tillage, irrigation, grazing and nomadism, plants
and animals, folk-lore and religion, monuments and art, politics and
industries, and carefully thought out geographical and historical data for
the serious student, Mr Rickmers's book is exceedingly attractive.

Westminster Gazette. — Mr Rickmers's book is pleasantly seasoned with the
incidents of Eastern travel, with accounts of climbs which will appeal to
mountaineers, and many a picturesque description of the singular but
imposing scenery. On the other hand, it is a most careful and scientific
study, illustrated on every other page with excellent and well-chosen
photographs of the physiography of this region.

Saturday Review. — A book, eloquent, enthusiastic, and learned, grandly illus-
trated, complete with scientific essays, glossary, bibliography, and a subject
index that indicates the immense scope of his observations and interests.

Mr Rickmers's broad elucidation, during the course of the narrative,

of the physical geography of the Duab and its Asian significance is
masterly writing that will repay the attention of others besides critical
scholars. Apart too from this main scientific inquiry, Mr Rickmers has
plenty to say that is more than merely interesting and amusing to the
general reader, for he conveys from the very first pages an impression of
sound judgment and also his feeling for the romance of travel.

Outlook. — Although much of the book is devoted to the physiography of the
Duab, it is also and always a delightful description of travel, and the

general reader may be assured of agreeable entertainment A notable

feature of the book is its superb illustrations, which, though reproductions
of photographs, are astonishingly artistic in treatment.

\_A special 8 //. prospectus, with specimen pages and illustrations,
will be foni'arded on application^

CAMBRIDGE

MANUALS OF SCIENCE AND
LITERATURE

Editors: P. Giles, Litt.D., Master of Emmanuel College
A. C. Seward, M.A., F.R.S., Professor of Botany
in the University of Cambridge

Price : is. net in cloth ; 2s. 6d. net in lambskin.

The following volumes will be ready shortly : —

Beyond the Atom. By Prof. J. Cox, M.A.
Bees and Wasps. By O. H. Latter, M.A.
The Wanderings of Animals. By H. F. Gadow, F.R.S.
Submerged Forests. By Clement Reid, F.R.S.
Wireles Telegraphy. By Prof. C. L. Foi'tescne, M.A.
The Theory of Money. By D. A. Barker, I.C.S.
English Monasteries. By A. Hamilton Thompson, M.A.
Mysticism in English Literature. By Miss C. F. E.

Spurgeon
Plato: Moral and Political Ideals. By Mrs A. M.

Adam.
Copartnership in Industry. By C. R. Fay, M.A.

The following volumes have already been published : —
History and Archaeology. Literary History.

Ancient Assyria. By Rev. C. H. IV. Early Religions Poetry of tlu Hebrews.
Johns. By the Rei'. E. G. King.

Ancient Babylonia. By Rei\ C. H. W.

Johns.
A History of Civilisation in Palestine.

By Prof R. A. S. Macalistcr.
China and the Manchus. By Prof

H. A. Giles.
The Civilization of Ancient Mexico.

By Lewis Spence.
The Vikings. By Prof. Allen Mawer.
New Zealand. By the Hon. Sir

Robert and J. Logan Stout.
The Ground Plan and the Historical

Gro7vth of the English Parish

Church. {2 vols.) By A. Hamilton

Thompson.
Brasses. By J. S. M. Ward.
Ancient Stained and Painted Glass.

Bv F. Svdnev Eden.

Early Religious Poetry of Persia. By

the Rev. Prof J. II. Moulton.
The History of the English Bible.

By the Rev. John Brown.
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CONTENTS— Continued

Mathematics and Physics

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Bibliography

Catalogue of the MSS in the Library of Corpus Christi College,
Part VII

Cambridge University

Christ's College Register, Vol. II

Biographical History of Gonville and Caius College

Scientific and Literary Journals

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The Journal of Hygiene Plague Supplement 11

Parasitology . .

The Journal of Physiology

The Journal of Genetics .........

The Biochemical Journal

Biometrika

The British Journal of Psychology .......

The Journal of Agricultural Science

page

15

16

17
17
26
26

19
20
21

18
27

27

23

27
26

27

27
28

28

28
28
28
28
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28

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Volume III

SEPTExMBER, 1913

No. 2

REDUPLICATION SERIES IN SWEET PEAS.

"""Tunica'.

By R. C. PUNNETT, M.A., F.R.S.

CONTENTS.

PAGE

Introduction . 77

A. The B, E, L series :

(1) The relation between B and L 78

(2) „ „ „ B and E 80

(3) „ „ ,, E and L 80

(4) The relations between B, E, and L in plants heterozygous

for all three factors 81

B. The D, F, N series :

(1) The relation between D and F 84

(2) „ „ „ F and N Si

(3) „ ,, „ N and D 88

(4) The relations between D, F, and N in plants heterozygous

for all three factors . 88

Primary and secondary reduplications 90

Conclusion 93

Literature ............. 95

Tables 96

Introduction.

The present paper aims at bringing together some of the results of
work done with sweet peas during the seasons 1908-13, and especially
that part of it which concerns the phenomenon of the reduplication of
terms in the gametic series.' Some of the results have already been
published in two short papers (Bateson and Punnett, (4) and (5))\
The work is far from completion, but as the subject is beginning to
excite greater interest it has seemed advisable to collect together and
publish these records for the use of other students. Many new points

' These figui'es refer to the list of literature at the end of this paper.
Journ, of Gen. iii G

78 Redvpliratkm Series in Street Peas

have arisen, especially (luring the present summer, and several years
must necessarily elapse before they can be settled satisfactorily.

The work recorded in the follnwing pages deals with the inheritance
of six factors, viz. :

B the factor for blue as opposed to red.

E „ „ erect standard as opposed to hooded.

L „ „ long pollen „ „ round.

D „ „ dark axil „ „ light axil.

F „ „ fertile anthers „ „ sterile.

N „ „ normal flower „ „ cretin.

As pointed out elsewhere ((4), p. 4) the relation between the factors
B, E, and L is such that a plant which is heterozygous for any pair of
this series shews coupling or repulsion according as the mating is
AB X ab, or Ab x aB. Until the present summer (1913) the experi-
mental evidence was in favour of regarding this generalisation as true
also for the series D, F, N, and to some extent it certainly is true. But,
as will appear later, there are grounds for supposing that under certain
conditions, hitherto undetermined, a simple Mendelian relation, without
coupling or repulsion, is to be found for this trio of factors. In this
connection it is interesting to recall Baui's work with Antirrliinam
(1) suggesting that in this genus coupling may result from the cross
AB X ab, while for the same pair of characters the mating Ab x aB
may give an ordinary Mendelian result. In considering the data given
in the present paper, those relating to the B, E, L series will be taken
first.

A. The B, E, L Series.

(1) The relatiim between B (ind L in plants huntozyguus fvr

E or e.

(a) Nature of mating BL x bl.

This was the reduplication series first discovered, and the nature
of the F„ generation can be explained on the hypothesis that F^
plants made in this way produce a series of gametes of the form
7BL : IBI : IbL : 7bl. To the evidence already given, derived from F.,
families ex Emily Henderson long pollen x E. H. round pollen ((2),
p. 36), can now be added 4 further F^ families (Table I, A), 40 F-^
families (Table I, B), and 14 families (Table I, C) derived from various
crosses inside the E. H. strain. The entire material was homozygous

R. C. PUNNKTT 79

for E. As Table I shews the results from these three sources are all
in accordance and confirm the original view that the results are probably
the outcome of a gametic series of the form 7BL : IBI : IbL : 7bl.
That the reduplication is of the same form in both ^ and % gametes
is supported by some experiments made in pollinating red round-
pollened plants with F-^ plants ex BL x bl. 112 jjlants were raised in
this way and consisted of

Purple long 50, Expectation being 1^9.

,, round 7, „ „ 7.

Red long 8, „ „ 7.

round 47, „ „ ^P.

The figures make it evident that the ^ gametic series is of the
form suggested, and as the F„ results can only be explained on the
supposition that this series of ^ gametes fertilizes a similar series of
% gametes there is every reason for supposing that the series is of the
same form in both. This will be assumed throughout the various
cases discussed.

It is assumed therefore that the reduplication between B and L is
on the 7 : 1 basis inside the E. H. strain, and though in some families
the two forms Bl, bL are in excess while in others they are deficient
these will be regarded as chance results. Further there is some evi-
dence to shew that the nature of the reduplication may remain
unchanged when the E. H. strain is crossed with some other strain.
Eight such families are given in Table II and shew that the expec-
tation on the 7 : 1 basis is closely realised. These families also afford
evidence that the form of the colour-pollen reduplication is unaf-
fected by the presence of sterility in the families.

There are however other cases in which the reduplication departs
considerably from the 7 : 1 basis. These are families derived from
plants which are also heterozygous for E. To these we shall return
later (p. 81).

(/3) Nature of mating Bl x bL.

Four families containing 419 plants were raised from this form of
mating and the numbers of the four classes were

Purple long Purple roun<l Red long Red round

226 9.5 97 1

It has been pointed out elsewhere ((5), p. 297) that this result is
in harmony with the supposition that the gametic series was

IBL : 7BI ; 7bL : Ibl.

6-2

so Brdifj)Iicafiou Scries in Suieet Peas

The four feiuilies consisted of hooded plants only and were not of the
pure E. H. strain. One small family however was raised from a cross
between a blue round and a red long in the E. H. strain. The F«
('11, 43) consisted of 8 blue long, 3 blue round, and 5 red long and
there is little doubt that if larger numbers were available the series
would resemble that just dealt with.

(2) The relation between B mid E //( families homozijrjous

for L or I.

(a) Nature of mating BE x be.

25 families have been raised from plants of this type of mating
and as is shewn in Table III the distribution of the four classes
among the 2712 jJants accords well with the view that the gametic
series is 127BE : IBe : IbE : 127be. All the plants were homozygous
in L.

(/3) Nature of mating Be x bE.

Many of the families in which this form of mating occurred were
also heterozygous in pollen, and these will be found recorded in
Table VI. The result however was the same throughout. No red
hooded jilant was evoi- found, and the other three classes occur approxi-
mately in the ratio 2 blue erect : 1 blue hood : 1 red erect. The total
numbers (including those in Table VI) are

Blue erect Blue linod Red erect

2969 1379 1441

This result points to the Be and bE types of gamete being con-
siderably more numerous than the other two, and it is not incompatible
with the view that the series is IBE : 127Be : 127bE : Ibe. But as
only one hooded red is to be looked for in 65,536 plants it is not pro-
posed to investigate this ]]articular case any further.

(3) The relation between E and L in plaids honiozyrjons
for B or b.

(a) Nature of mating EL x el.

At present there are no records of such a cross though a number of
such families are expected in 1914.

(y8) Nature of mating El x eL.

Four such families were raised in 1908 and as is shewn in Table IV

E. a PUNNETT 81

they consisted of the three classes erect long, erect round, and hooded
long approximately in the ratio 2:1:1. It is probable that the rela-
tion between E and L is similar to that existing between B and L, and
between B and E, but at present the exact nature of the series must
be left undecided.

(4) The reldtioiis between B, E, and L in plants tvhich ure
heterozygous in all three factors.

As previously pointed out ((4), p. 7) there are four ways of mating
to produce such plants, viz. ;

(a) BEL X bel,

{13) BeL X bEI,

(7) BEI X beL,

(S) bEL X Bel.

Owing to misadventure an F., generation has been raised from the
first two crosses only and we may now proceed to consider the results
separately.

(a) Nature of mating BEL x bel.

The appearance of a single red hooded round plant in 1910 (cf (5),
p. 297) made this mating possible and an F., generation was raised in
1912. Details of 13 such families are given in Table V. The results
are evidently complex but some attempt at analysis may be made by
considering separately each pair of the three factors concerned. The
figures then become

(«)

BL : Bl

bL

bl .

: 479 : .58

66

143

ih)

BE : Be

bE

be:

: 532 : 5

6

203

(c)

EL : El

. sL

el :

: 479 : .59

66

142

In each case the nature of the gametic series is evidently « : 1 : 1 : ?(
where n > 1. Since for any two of the factors the nature of the
original cross was AB x ab the " coupling " form of the series in each
case was to be expected. But it is equally evident that the value of n
in cases (a) and (6) is different from what it would have been had the
plants been heterozygous for the two factors in question alone. The
experimental numbers in (a) are not on the 7 : 1 system but the results
are given far more closely by the gametic series 5BL : IBI : IbL : 5bl.
On this supposition expectation would be 501 blue long : 57 bhn; red :

82

Bedv pi! cation Series in Sweet Peas

57 red long : 12!) red round — iiuinbers not far removed from those
actually found. The value of ii for the EL series must also be very
close to its value in the BL series.

Again in the BE series the figures obtained are much more closely
in accordance with the supposition tiiat the value of /( is 08 instead of
127 which was found to be the case where the plants were homozygous
for L (cf. p. 80). Apparently what may be termed the normal linkage
ratios are upset, but this point will be considered later (p. 91).

It may however be suggested that it is possible theoretically to
construct a gametic series which would give a close approach to the
figures obtained. Such a series is :

103 BEL
23 BEI
1 BeL
1 Bel
1 bEI
1 bEL
23 beL
103 be I

In such a series the gametic relations for the separate pairs of
factors are :

B and L— 13BL : 3BI : 3bL : 13bl,

B and E— (i3BE : IBe : IbE : 63be,

L and E— 13LE : 3Le : 3IE : 13le.

As the subjoined figures shew the theoretical results accord fairly
closely with the actual numbers.

Blue Red

giving the zygotic figures in
the pi'opoi'tion

BEL 43283

BEI

.5361

BeL

310

Bel

207

bEI

207

bEL

301

beL

5267

be!

10609

Erect Hood

Long Round Long Round
477 55 2 3

Erect

Long Round Long Round
•2 4 64 139

61

a-5

3-5

2-5

60

120

Expectation 4!)3

At present however I do not wish to lay stress on this approxima-
tion as I am unable to picture clearly how such a gametic series comes
to be formed.

(/8) Nature of mating BeL x bEI.

Details of a number of F., and F^ families ex Blanche Burjiee

R. C. PUNNETT 83

(hooded white long) x E. H. louiid were given in Report IV, Evol.
Gomm. Roy. Soc. 1908, p. 14. The general result shewed "coupling"
between B and L, together with " repulsion " between B and E, and
between L and E, a result which from the nature of the mating was to
have been expected. It was pointed out in an earlier account (R. E. G.
IV. p. 11), that the two classes purple round and red long were less
numerous than was expected on a 7 : 1 basis, and at the time an ex-
planation was suggested on the grounds of some of the families being
on a 7 : 1 basis and others on a 15 : 1 basis. The recent publication of
Trow's Paper (16) on reduplication series has put the matter in a new
light and we shall return to this case later in discussing his suggestion
(p. 91). For the moment we may turn to Table VI which adds further
data by the inclusion of a number of t\ and F3 families derived from
the cross white hooded Bush long (BeL) x white erect Cupid round
(bEI). The F^ families were given in the earlier account ((3), p. 12)
and were there regarded as a case of coupling on a 15 : 1 basis. Fuller
experience however has led to the conclusion that they should be classed
with the material derived from the original cross Blanche Burpee x
Emily Henderson. Three further families ('08, 89, 93, 114) have
been added derived from other material in which the mating was of
the nature BeL x bEI.

Consideration of each pair of the three factors taken separately
shews their relations to be as follows :

(a) BL : Bl : bL : bl :: 3006 : 164 : 212 : 843,

(^) BE : Be : bE : be :: 2146 : 1024 : 1055 : —

(7) EL : El : eL : el :: 2200 : 1001 : 1018 : 6.

Evidently (yS) and (7) are " repulsion " forms of reduplication in
which the zygotic series 2n- + lAB : ?!- — lAb : )z'-'— laB : lab is de-
rived from a gametic series lAB : {n — l)Ab : (n — l)aB : lab ((5),
p. 295). An approximation to the value of ?( in such cases is most
readily obtained by dividing the last term in the zygotic series into
one of the two middle terms. This gives the approximate value of
n- — 1 from which the value of n may be readily deduced.

In the case of (7) — - — = — — -^ whence n- — 1 = 169 and « = ap-
proximately 13. Hence the zygotic series (7) is given most nearly by
a gametic series lEL : 12EI : 12eL : lei.

In (/S) there is no individual lacking both B and E, and all that can
be stated of n is that it is almost certainly > 32.

84 Reduplicafion Scries /» Suieet Peas

The zygotic series (a) is evidently a "coupling" form of reduplica-
tion, and is most nearly approached by the gametic series 10:1:1:10,
on which the expectation of the 4 forms is 2984 : 184 : 184 : 873. It
is clear that in this case, as in the BEL and bel cross, the normal value
of the pollen-colour reduplication series is upset, and that instead of
being on a 7:1 basis it approximates to a 10 : 1 basis. It is possible
that this upsetting of the normal value may be true also of the
standard-pollen, and of the standard-colour series, but at present the
point must remain undecided. For we do not know the normal value
of the standard-pollen series, and in the case of the standard-colour
series the data are insufficient for determining even its approximate
value.

B. The D, F, N Series.
(1) The relation between D and F in plants homozygous for N.

(a) Nature of mating FD x fd.

Evidence has already been given ((3), p. 16) for regarding the re-
duplication in this case as on a 15 : 1 basis, and to the figures previously
.set out we can now add a few more families (cf Table VII). As the
figures shew, the numbers of the two classes, dark sterile and light
fertile, are rather below expectation. Certain of the families (marked
* in Table VII) were also heterozygous for B and L and it is po.ssible
that this may have infiuenced the result. Certainly when these families
are deducted the discrepancy in the remainder between reality and
expectation is not so marked (cf Table VII).

(/3) Nature of mating Fd x f D.

The experimental data for this mating are given in Table VIII.
So far no light axilled sterile has occurred among 2252 plants, though
2 such plants were to be expected if the gametic series is on the
1:15 basis. At present therefore the value of the reduplication is un-
decided.

(2) The relation between F and N.

Details of fl F., families derived from the mating Nf x nF have
been given elsewhere ((5), p. 295) and it was clear that the nature of
the gametic series produced by the F^ plant was INF : .3Nf : 3nF : Inf.
In the same paper it was suggested that the value of the reduplication
affecting two factors miglit be the same for both the "coupling" and

R. C. PUNNETT 85

the "repulsion" series, and the low value of n in the present case
offered the most favourable opportunity hitherto met with for testing
this point. Accordingly large numbers of pollinations from normal
fertile plants (NF) were made on to sterile cretins (nf). In not a single
instance however was the operation successful and it was found subse-
quently that the cretin, whether producing pollen or not, is always
sterile on the female side. The pods, and even the seeds, may some-
times undergo swelling but in none of the numerous cases examined
was a viable seed produced. The (NF x nf) plant had therefore to be
sought for in another direction.

Since the gametic series produced by the plant formed from the
two gametes Nf x nF is of the nature INF : ."^.Nf : 8nF : Inf such
plants should give rise to the following zygotic forms :

NNFF 1 NNff...9 nnFF ... 9 nnff ... 1

NNFf 6 Nnff ... 6 nnFf ... 6

NnFF 6

(Nf X nF) ... 18
(NF X nf) ... 2

normal] cretinl cretin]

normal fertile SS : , ., ^ 15 : „ ., } lo : , ., ^ 1

sterile ) fertile] sterile)

Hence out of 33 normal fertiles in such a family there should be 5
distinct classes distributed in the following proportions :

1 (NNFF) giving only normal fertiles.

6 (NNFf) giving normal fertiles and steriles only.

6 (NnFF) giving normal and cretin fertiles only.

18 (Nf X nF) giving all 4 classes with "repulsion" between N
and F.

2 (NF X nf) giving all 4 classes with "coupling" between N
and F.

30 such normal fertiles from F., families were grown on to give an
Fa generation in 1912. The expected 5 classes were all found, and as
shewn below (and also in Table IX) the proportions in which they
occurred accorded closely with expectation.

86 Reduplication Series in Sweet. Peas

Type of -A'.j plant

Number found
hy experiment

Number
expected

NNFF

2

•9

NNFf

0

5-5

NnFF ...

7

.5 -.5

(Nf X nF)

14

16-3

(NFx nf)

2

1-8

Totals ... 30 30

In one family where coupling occurred ('12, 76) the numbers were
sufficiently large to indicate the nature of the reduplication. There is
little doubt that it is on the 3 : 1 basis for although there was an
unexpectedly large number of cretins the proportions accord fairly well
with expectation'.

Normal NornLil Cretin Cretin

fertile sterile fertile sterile

12; 76 90 20 30 34

Expectation 111-5 19 19 34-S

It should be mentioned that the plants in this family were all light-
axilled. There are good grounds then for asserting that for the factors
N and F reduplication is on the 3:1 or on the 1 : 3 basis according as
to whether the original mating is NF x nf or Nf x nF.

Yet it is probable that the case is not so simple as it appears at first
sight. Refei-ence to Table IX shews that in two families, viz. '12, 71
and '12, 91, the proportions in which the four forms occur afford a
distinct suggestion of a 9:3:3:1 ratio. And if these two families
are taken together the figures 42 NF, 13 Nf, 14 nF, 5 nf make a very
close approximation to this ratio.

The jjossibility that, where the factors N and F are concerned, there
may be other sorts of families than those shewing the 3 : 1 or the 1 : 3
reduplications is to some extent supported by the results of growing
on a further generation from the F-j fixmily '12, 76. Unfortunately
these did not set seed freely and only 11 F^ families were obtained.
In all of them the number of individuals was relatively small, and in
two cases it was too small to give more than a qualitative result.
There are however 8 families in which all the four classes occurred,
and in which there are over 30 plants jjer family. Reference to
Table X shews that these families' fall into two distinct groups : —

' I do not feel clear as to the nature of Fam. '13, 47 and have therefore not included
it in the present argument. It may be a family on the 3:1:1:3 basis but the numbers
are too few for certainty.

R. C. PUNNETT 87

(a) Fanis. 38, 39, 40, 41. These four families are evidently of the
same nature as the parent family '12, 76. The three classes
normal sterile, cretin fertile, and cretin sterile are much in
the iDroportions that would be expected on a 3:1:1:3 re-
duplication system, but the number of normal fertiles is much
below expectation. These four families added to the parent
family ('12, 76) give the numbers :

Normal Normal Cretin Cretin

fertile sterile fertile sterile

165 58 58 78

Expect, on 3 : 1 : 1 : 3 basis ... 346 59 59 76

The italicised figures shew that the three last classes are closely in
the ratio 7 : 7 : 9, as expected on the 3:1:1:3 system, but that the
normal fertiles are only about one half as numerous as would be ex-
pected. That there is something militating against the formation of
normal fertiles in such families is borne out by the following considera-
tion. Of the 11 Fi families grown from '12, 76, 10 contained all four
classes of plant, while one family had cretins but no steriles. Now
the normal fertiles from a family producing gametes on the system
3 NF : 1 Nf : 1 nF : 3 nf should be of five kinds, and in every 41 plants
these five kinds should be present in the following proportions:

9 NNFF giving normal fertiles only.

6 N N Ff ,, normal fertiles and steriles only.

6 NnFF ,, normal and cretin fertiles only.

18 NnFf ,, all four classes with coupling.

2 NnFf „ „ „ „ repulsion.

Hence out of 41 such plants only 20, or not quite one half, should give
the four classes. But of the 11 plants tested (Table X) no less than
10 gave all four classes. It would appear then that for some reason
or other the classes NNFF, NNFf, NnFF are not produced in the ex-
pected numbers, though why this should be so is at present quite
obscure.

(h) Fams. 43, 45, 46. These three families are distinctly different
from those just considered and the combined figures

Normal Normal Cretin Cretin

fertile sterile fertile sterile

75 23 21 5

Expect 69-75 23-25 23-25 7-75

88 EedtipEcafion Series in Sweet Peas

support the idea that we are here dealing with a 9:3:3:1
ratio. As ah'eady menticined it is not impossible that in
Fam. '12, 71 (Table IX) this type of femily has been encoun-
tered once before.

(3) The relation between N (iiid D in plants Immozygous for F.

Hitherto the opportunity of studying this particular case has been
limited to a single family. This family (13, 119) was from a normal
fertile belonging to Fam. 12, 88 (Table IX)' and proved to be hetero-
zygous in N and D but homozygous f(jr F. It contained 100 plants
and was composed of

Normal Normal Cretin Cretin

dark light dark light

48 22 27 3

Expect, on 1 :3 :3 : 1 system ... 51-6 23-i 23-i 1-6

The figures are in consonance with the idea that the reduplication
for N and D is on a 1 : 3 basis, and this is borne out by the behaviour
of these two factors in relation to one another among plants which are
heterozygous for N, D, and F (see p. 89).

(4) The relations between D, F, and N in plants heterozygous

for all three factors.

As in the case of the B, E, L series there are four ways theoretically
possible by which plants heterozygous for three factors can be pro-
duced, viz. :

(a) DFN X dfn.

(fi) DfNxdFn.

(7) DFnxdfN.

(h) dFN X Dfn.

Of these four ways I have hitherto been able to study two, viz. (13)
and (7).

(/3) Nature of viatinr/, DfN x dFn.

The details of four such families raised from the same parents and
comprising in all 442 plants are set out in Table XI, and may be
considered together. The proportion of dark axils to light a.xils, of
fertilea to steriles, and of normals to cretins is closely 3:1 in each case.

R. C. PuNiNETT 89

And if the figures are arranged for the three different pairs of factors
the result is what might have been anticipated from the nature of the
mating, viz. coupling between D and N,and repulsion between D and F
and between N and F respectively,

Families

48—51.

Expectation on
3:1:1:3 system.

ND

282

^84

Nd

49

48

nD

52

48

nd

59

62

Expectation on
1:3:3:1 system.

NF

225

228

Nf

106

103:5

nF

101

103:5

nf

10

7

Expectation on
1:7:7:1 system.

Expectation on
1 : 15 ; 15 : 1 syster

DF

220

222'7

221-4

Df

114

108-8

110-1

dF

106

108-8

110-1

df

2

1-7

-4

The numbers shew that the reduplication series in the ,first two
cases is approximately on a 3 : 1 basis while that for D and F fits most
closely a 1 : 7 : 7 : 1 series. In the last case however the df class appears
so rarely that the precise nature of the reduplication must for the
present remain undecided. It is not impossible that it may be on a
1:15 basis which is rather to be expected in view of the fact that the
coupling series for these two factors is on the 15 : 1 : 1 : 15 system.

It may be noted that the 3 : 1 basis for the relation between N and
D from this mating is in accordance with the 1 : 3 basis for the relation
between these two factors when the natui-e of the mating is Nd x nD.

(7) Nature of mating, DFn x dfN.

The material for the study of this mating (Table XI) consists
so far of two F^ families, '13, 52 and '13, 53, and of four F^ families,
'13, 113, 114, 117, 123, all derived from an original cross of the same
type. All six families appear to be of the same sort, and I have
assumed the nature of the mating in the F^ families is what is known
to be the case in the two F„ families.

In families derived from this type of mating I had anticipated

9ft RedupUcaftoii Series lii Sweet Peas

coupling between D and F, together with repulsion between N and F
and between N and D. The figures here given shew that the first ])art
of the anticipation was realised.

KamUies 32, 53.
IH. 117. 123.

n3.

Expectation on
15 : 1 : 1 : 15 system.

DF

17'2

170-4

Df

6

7-3

dF

4

7-3

df

68

55-0

There is coupling between D and F and this is not far removed from
the expected 15 : 1 : 1 : 15 system. But there is no repulsion evident
between either N and F, or between N and D.

Families 52. 53, 113,
IH, 117. 123

Expectation.

ND

ni

135

Nd

49

45

nD

47

45

nd

13

15

NF

l.?0

135

Nf

50

45

nF

46

45

nf

14

15

Instead of this the figures accord closely in both cases with an
ordinary 9:3:3:1 ratio. The possibility of a similar ratio has already
been referred to in the case of other families where these factors are
concerned (p. 86) and it is hoped that a further study of this interest-
ing case may give us the clue as to the relation between normal and
reduplicated gametic series. But it is evident that several years must
elapse before the necessary experiments can be completed.

Primary and Secondary Redui'Lication.s.

In an interesting paper which has just appeared (16) Trow dis-
cusses the problems offered by reduplicated series of gametes, and
draws a distinction between what he has termed primary and secondary
series. Starting with the case where there are three factors A, B, C
such that any two may form a reduplicated series, he has shewn theo-
retically that if the reduplication between A and B is of the form
III : 1 : 1 : m, and between A and C is of the form n : I : 1 : ii, then
the secondary form of reduplication, derived from these two primary
ones and expressing the relation between B and C, is of the form
iiiii+ 1 : lit +11 : 111+ n -.11111+1. The only experimental data with

R. C. PUNNETT 01

which he has hitherto been able to test his hypothesis have been pro-
vided by a case among Gregory's primulas, the three factors concerned
being M (= magenta), S(= short style), and G(= green stigma). The
experimental evidence points to the MS reduplication being on the
7 : 1 basis and the MG reduplication on the 2 : 1 basis. The secondary
reduplication between S and G should consequently be of the form
(7 X 2) + 1 : 7 + 2 : 7 + 2 : (7 X 2) + 1, i.e. 5:3:3: 5, and the experi-
mental data are in fairly close accordance with this expectation (cf
Trow (16), p. 316). The data given in this paper from the two sweet
pea crosses BEL x bel, and BeL x bEI aftbrd further cases for testing
Trow's hypothesis.

(1) The BEL X bel cross.

It has been pointed out above (p. 82) that the three reduplication
series here are approximately of the following values :

(a) B and L— 13 : 3 : 3 : 13,
(/5) B and E— 63 : 1 : 1 : 63,
(7) E and L— 13 : 3 : 3 : 13.

If we consider (a) and (/S) as the two primary redup)lications then
the value of (7) should be

(J^i X 63) + 1 : J^'- + 63 : if + 63 : (J^ x 63) -I- 1,

i.e. 4-1:1 : 1 : 4-1,

a result which is closely in accordance with the experimental numbers.
A feature of interest in connection with thi.s case is that it demonstrates
that when the value of one primary series is considerably greater than
the other, the value of the secondary series will be very close to that of
the lower primary series.

(2) The BeL x bEI cross.

In this case the relations between B and E, and between L and E,
are of the nature of " repulsion." Nevertheless, as Trow has pointed
out, even if these be regarded as the two primary series the secondary
series (between B and L here) should be of the nature of "coupling."
The facts (p. 83) point to the reduplication between L and E being
near 1 : 12 : 12 : 1. That between B and E is evidently much higher
though how much so we have no direct means of telling. But we have
already seen that the "coupling" between B and E is on the 127 : 1
basis in families homozygous for L and on the 63 : 1 basis in the

92 Rediqilieatioii Series in Sweet Peas

BEL X be I cross. Also there is evidence for supposing that the
"repulsion" reduplication is of the same value as the "coupling" re-
duplication between two given factors. We shall assume then for
argument that the reduplication between B and E is of the value
1 : 63 : 63 : 1. For B and L (p. 84) the experimental data are closely
approached by a series 10 : 1 : 1 : 10. We have therefore the 3 series :

(c() B and E— 1 : 63 : 63 : 1,
(/3) E and L^ 1 : 12 : 12 : 1,
(7) B and L— 10 : 1 : 1 : 10.

If we treat (a) and (/S) as the primary series the value of (7)
deduced theoretically should be

(63 X 12) + 1 : 63 + 12 : 63 + 12 : (63 x 12) + 1

i.e. 1001 : 1 : 1 : lO'Ol,

which is a remarkably close approximation to the experimental pro-
portions.

It will be noticed that in each of the above cases we have chosen
as our two primary .series those in which the reduplication values are
highest. This was also done by Trow in his analysis of Gregory's
primulas. It is only in this way that the hypothesis will work, for, as
can be readily shewn, the value of the redujjlication in the secondary
series must always be less than in either of the two primary series
from which it is derived.

Let A, B, and C be the three factors concerned, and let the redupli-
cation series for

A and B = p : 1 : 1 : p (a),

A and C=p-\-x:\ -.1 : p + .c (/S),

where /; > 1 and x is positive.

Then the series for

B and C=p{p+ x) -1-1 : 2p 4- .r : 'Ip + ,/• : p (p + ./) 4-1 (7)-

It is required to shew that

2)jj} + x)j^l p

2p + .v r

i.e. p" + px -i- I < 2/r +j)x,

i.e. 1 < p".

which is evident since on hypothesis /) > 1.

R C. PUNKETT 93

Hence in a series of three reduplications, two primary and one
secondary, that one is to be regarded as the secondary in which the value
of the reduplication is lowest.

In this connection the two groups of families recorded in Table XI
are also of interest. In Group A from the mating DfN x dFn there is
repulsion between D and F, coupling between D antl N, and repulsion
between N and F. The first repulsion is not improbably on the 1 :15:15:1
system or something near it (cf j). 89), while the second repulsion and
the coupling are not far removed fi-om the 1:3:3:1 and the 3:1:1:3
systems respectively. The figvu-es are in general accordance with Trow's
hypothesis, but tlie numbers are not large enough to determine more
precisely the values of the reduplication systems, or tci decide which of
the two lower .series is the secondary one.

In the families of Group B in Table XI, where the nature of the
mating is dfN x DFn, there is coupling between D and F but no
repulsion between N and D or between N and F. Here again the
experimental facts are in accordance with Trow's hypothesis, for where
one of the primary series shews no reduplication it follows that no
reduplication will be exhibited by the secondary series.

*

Conclusion.

Finally attention may be drawn to some points in connection with
the value of the reduplication in the various cases discussed above.
Where only two factors are concerned we have regarded the redupli-
cation as of the form (?; — 1) : 1 where n is some power of 2, and we
suggested in a previous paper how such a series might be brought
about through alternating periclinal and anticlinal cell divisions ((5),
Fig. 4). The experimental data hitherto obtained from sweet peas fit
in with this view, but, as pointed out some years ago ((2), p. 9), they
are also in accordance with the form of the reduplication being
n : 1 : 1 : « where n again is some power of 2. It is only in cases
where n is very small that we can hope to distinguish between the
two without growing an impracticably large number of plants. At
present the NF x nf mating is the only one from which we can look
for a critical result on this point, and the available evidence suggests
that the reduplication here is 3NF : INf : InF : 3nf (cf p. 86), i.e. that
it is on the {n — 1) : 1 basis rather than on the n : 1 basis'. It may

1 What is evidently a case of reJuplication on the 3 : 1 basis has been recently dis-
covered by Collins in maize ((6), p. 579), the cross in (juestion being one between an
Journ. of Gen. iii • 7

94

HediipUcation Series in Sweet Peas

be pointed out however that this scheme is not incompatible with the
)i : 1 basis. The form of the series, whether (71- 1) : 1 or n : 1, might
depend upon whether the first division in the quadrant were a periclinal
or an anticlinal division. In the one case (Fig. 1) we should get the
(k- 1) : 1 : 1 : (w- 1) series, and in the second case (Fig. 2) the

2nd division

1st dirisiun

1st (lirisioii
: 1st divisiuii

'Jnd dii'isiuyi,

Fig. 1.

Fig. 2.

?i : 1 : 1 : n series. Further possible ratios also depend upon whether
the first division in the other two quadrants is periclinal or anticlinal.
Indeed it is obvious that there are numerous possibilities which may
perhaps repay discussion when more experimental data are available.
All that can be stated positively at present is that the cases hitherto
worked out in the sweet pea fit in with the hypothesis that the number
of cells in the reduplicated series is some powci- of 2 where only two
factors are concerned. But where three factors are concerned this is
certainly not true. The value of the primary reduplications is evi-
dently altered, and there would seem to be some process whereby these
reduplications react upon one another. Where so many points remain

American variety with coloured aleurone and horny endosperm, and a Chinese variety with
white aleurone and waxy endosperm. By means of other statistics Collins is at pains to
prove that the reduplication phenomena in maize are of a hifjlily irregular nature. Much
stress however cannot be laid upon these results as the author is evidently dealing with
dominant as well as recessive whites in his experiments though this point does not appear
to be specifically recognised by him. It is probable that a more careful genetic analysis
of the whites which he uses would help to clear up tlie apparent irregularities in his
results.

R. C. PUNNETT 95

doubtful, as at present, it is difficult to suggest any scheme by which
this result could be brought about, and the problem must at present
be left in the hope that fresh data may eventually lead to its solution.

LITERATURE.

(Papers dealing with cases in wliich the phenomena of "coupling'' and "repulsion"
are met with, but in which sex-limited inheritance is not involved.)

1. Bade, E. "Vererbungs- und Bastardierungsversuche mit Antirrhinum;

II. Faktorenkoppelung." Zeit. f. ind. Abst. u. Vererb. 1912.

2. B.iTESON, AV., Saunders, E. E., and Pdxnett, R. C. "Experimental Studies

in the Physiology of Heredity." Report III, EvoL Coimn. Roy. Soc.
1906.

3. . Report IV, Evol. Comm. Roy. Soc. 1908.

4. Bateson, W., and Pdnnett, R, C. "On the Inter-relation of Genetic

Factors." Proc. Roy. Soc. 1911.

5. . " On Gametic Series involving Reduplication of certain Terms."

Journal of O emetics, 1911.

6. Collins, G. N. "Gametic Coupling as a cause of Correlations." Atiier.

Nat. 1912.

7. Emerson, R. A. " Genetic Correlation and Spurious Allelomorphism in

Maize." Twenty-fourth Ann. Rep. Nebraska Agric. Exp. Stat. 1911.

8. Gregory, R. P. " On Gametic Coupling and Repulsion in Primula sinensis.^'

Proc. Roy. Soc. 1911.

9. Hagedoorn, a. L. " The Genetic Factors in the Development of the House-

mouse, &c." Zeit. f. ind. Abst. tt. Vererb. 1912.

10. Keeble, F. W., and Pellew, C. "The Mode of Inheritance of Stature and

of Time of Flowering in Peas." Journal of Genetics, 1910.

11. Morgan, T. H. "Complete Linkage in the Second Chromo.some of the

Male of Drosophila." Science, 1912.

1 2. PuNNETT, R. C. " Inheritance of Coat-Colour in Rabbits." Journal of

Oenetics, 1912.

1 3. Saunders, E. R. " Further experiments on the Inheritance of ' Double-

ness' and other Characters in Stocks." Journal of Oenetics, 1911.

14. Sturtevant, a. H. "The Linear Arrangement of Six Sex-linked Factors

in Drosophila, as shown by their Mode of Association." Journ. Exp.
Zool. 191.3.

15. Trow, A. H. "On the Inheritance of Certain Characters in the Common

Groundsel, &c." Journal of Oenetics, 1912.

16. . "Forms of Reduplication; Primary and Secondary." Journal of

Oenetics, 1913.

1 7. DE ViLMORiN, P., and Bateson, AV. "A Case of Gametic Coupling in Pi-sum."
Proc. Roy. Soc. 1911.

7—2

96

RedupUcation Series in Sioeet Peas

TABLE I.

Families from Emily Henderson, heterozygous in colour and pollen.
A. t\ families ex E. H. long x E. H. round.

Reference
Number

Pari

k-

Pico tee

Red

Tinged

white

Wl

lite

Long Round

Long Round

Long

Round

Long

Round Long

Round

'08, 119

12

3

4 —

3

2

1

3

24

10

„ l'-20

22

7

9 —

2

0

—

4

34

13

'09, 6

33

2

9 —

3

11

—

5

37

10

'10, 58

62

4

21 2

7

15

—

6

73

16

Totals

129

16

43 2

15

34

1

18

168

49

Purple

Red

White

Long Round

Long

Round

Long

Round

Above 4

families

... 172

18

16

52

168

49

Report III, p. 36

... 1528

106

117

381

1190

394

Totals

... 1700

124

133

433

1358

443

Expectation among

colouif'd 165^

liO

140

iSS

B. Fj families heterozygous in colour and pollen.
Purple Red

White

Reference

Number

Long

Round

Long

Round

Long

Round

04, 152

46

4

2

10

30

4

'04 F, 82

40

4

1

9

38

10

„ 102

20

—

1

2

7

3

„ 105

47

4

8

8

39

17

„ 106

12

—

1

7

5

3

,, 128

17

—

—

4

—

—

'05, 280

31

4

3

8

18

4

„ 286

8

1

1

2

7

1

,, 291

13

3

—

8

8

3

„ 292

31

4

1

10

17

2

„ 293

10

—

1

5

11

3

„ 296

20

—

2

4

15

3

'05 F, 23

23

3

5

6

11

'06 F, 401

128

14

8

38

102

29

„ 402

42

7

1

12

14

3

„ 404

17

3

—

4

17

5

,, 409

48

9

7

16

—

„ 410

60

6

3

14

—

„ 406

74

6

5

39

31

15

,, 412

21

2

—

7

6

„ 413

79

7

5

23

21

6

R. C. PUNNETT

97

B. F., families heterozygous in colour and pollen (conthwed).

Purple

Red

E

eference
dumber

Long

Round

,

,

Long

Round

Long

Round

'OG F

, 417

50

5

4

7

28

3

, 425

17

1

3

7

2

3

, 428

69

5

1

18

39

18

, 431

28

1

2

7

21

8

, 432

15

3

3

0

19

5

, 433

27

1

2

5

7

1

, 437

64

2

3

16

—

—

, 440

H6

3

1

13

7

3

, 441

84

(i

5

27

75

19

, 444

44

1

4

9

32

12

, 448

23

4

2

5

9

2

, 449

32

2

6

5

16

3

, 450

23

—

—

4

—

—

, 451

21

1

1

11

7

—

, 452

20

1

—

6

4

1

, 453

30

—

1

8

24

11

, 455

47

6

2

14

9

5

, 462

26

1

—

11

12

6

, 463

37

1

1

10

12

4

To

ala ...

1480

125

96

425

Kx

oectatio}!

1-170

l:!f>

13.5

406

C. Families from various crosses inside the Emily Henderson strain.

Purple

White

Reference
Number

Nature of cross

Long

■' ,

Round

Long

Round

Long

Round

'06, 359

F-y ex Pic. long x E. H. round

41

2

3

20

19

3

„ 391

F.i ex Pur. long x E. H. round

96

9

7

28

36

7

'07, 164

F-> ex P. L. round x Pic. long

32

6

5

8

—

—

„ 165

?, »' »)

19

2

2

3

—

—

„ 166

tj 11 -1

16

—

3

9

—

—

„ 167

11 H 5)

18

1

3

3

—

—

,, 1G8

F> ex Bed round x Wh. long (ex Pur.

22

3

1

8

11

2

„ 170

)» ,» J)

12

2

4

12

7

1

„ 171

1, )» 1'

20

—

1

9

5

1

„ 172

», >» 11

18

2

1

5

4

—

„ 173

) > ,1 11

13

1

—

4

3

1

'OS, 121

F-i ex Pur. long x T. W. round

68

G

8

16

—

—

„ 122

U ,, ))

213

19

17

61

—

^

,, 125

F„ ex Pic. long x T. W. round

147

10

10

35

—

—

Totals

735

G3

65

221

Expectation

749

63

63

H09

98

Reduplication Series in Sweet Peas

Purple

Bed

Long

Round

Long

Round

Total from Fnmilips A

1700

124

133

433

B

UsO

1 5

96

425

0

735

03

65

221

Totals

3'.115

312

294

1079

Expectiition

3871

328

328

1073

TABLE IIT.

FamiliKS shewing rmipliiiff hetirppu pxr/i/e ami lht>. erect stamlanl.
(Nature of mating BE x be.)

Purple

Red

Reference

.

Number

Erect

Hood

Erect

Hood

•08,

84

139

—

2

44

'09,

2

67

1

—

22

•10,

36

6

—

—

2

,,

37

3

—

—

3

,,

38

0

—

—

2

, ,

39

17

—

—

6

,,

40

9

—

—

2

,,

41

11

—

—

2

,,

46

22

—

—

5

,,

55

158

—

1

56

•10 F,

1

206

—

2

37

>, 2

—3

178

4

—

45

,,

4

74

—

1

29

,,

5

59

—

—

17

,,

13

(;o

—

—

18

,,

14

138

1

—

36

,,

15

125

—

1

50

,,

18

57

—

—

11

,,

19

71

1

—

29

,,

20

62

1

■ —

21

,,

21

48

—

—

20

,,

22

171

2

—

63

,,

23

123

—

—

38

,,

24

i(;8

2

2

71

♦)

25

58

—

1

25

Totals ...

2036

12

10

654

Expectution on 1
197 . 1 ;,„«;,. ,

2023

11

11

667

R. C. FUNNETT

99

TABLE II.

Slfrility fami/if.i homozygous in axil heterozygous in pollen.

Purple

White

Reference
Number

Long

Round

Lontr

Rnunrl

I'uvijIi

J Red

Long

Round

sterile

U5, T.

L. 2

i'lo

11

8

9

12

7

48

17

20

,, ,,

5

75

9

7

13

19

7

48

23

20

'06,

340

■28

4

3

ir,

20

—

28

8

9

»)

3(30

33

3

3

11

15

3

31

9

10

'07,

90

154

11

6

40

45

23

54

15

22

■08,

981

r,3

4

7

15

22

i;

—

—

—

1*

100

(i4

't

6

18

29

11

IG

8

13

'09,

3

25

0

2

10

11

2

5

—

5

Totals

497

49

42

132

173

59

Kxpech
7: 1

ation on)
basis )

498

43

42

138

TABLE Y.

Families heterozygous in standard, colour, and pollen.
(Nature of mating EBL x ebl.)
Purple Red White

Reference
Nunil)er

Long

Eou;

'12,

21

20

3

22

71

7

23

CO

4

24

BO

7

26

58

4

27

20

2

28

13

1

30

34

4

31

14

4

32

4G

3

33

40

9

34

17

4

35

12

3

Erect
Long Round Long Round Long Round Long Round Long Round Long Round

1 —

7

10

—

—

—

—

y

19

20

—

2

5

10

16

20

1

3

5

3

20

—

—

-^

—

7

11

25

3

1

3

1

5

10

—

1

1

3

2

1

1

2

3

;i

14

6

1

1

1

3

9

19

—

—

4

4

14

6

2

3

3

5

14

31

4

3

1

1

4

9

3

1

5

2

1

—

—

—

1

Totals 477 55

64 139 147

15

17

32

' A sister plant of '08, 98, and '08, 100. gave a remarkably aberrant result. This was
'08, 99 in which there were 182 plants. Of these 180 were fertile and only 2 were sterile.
As in the sister plants the coloured were all deep purples and Miss Hunt and there is no
reason to suspect any error. This almost complete absence of steriles is quite unlike
anything else we have encountered and we are unable to offer any explanation.

100

RedHplication Series in Sweet Peas

TABLE IV.

Families slieiciny repuhion hflvwen erect slaiuJiinl and long pollen.
(Nature of mating El x eL.)

Red. Erect Ke<l- Hood

Reference
Number

Long

Round

Long

Round

'08, 9i

46

•21

30

—

„ 95

33

11

10

—

„ 90

46

15

16

—

„ 97

34

24

10

—

Totals

159

71

66

—

Expectation ...

US

7i

TABLE

VL

7i

Families he/eroxijgoii.t for pollen, colour, and hood.
(Nature of matiiiK BLe ■; blE.)
Purple Red

White

Erect

Hood

Reference
Number

Long

Round

Long

Round

Long

Round

Long

Round

Long

Round

^ /

■07,

80

15

•2

3

—

—

10

12

3

9

—

E

X

,,

81

17

—

13

—

3

10

15

3

4

—

,,

82

6

1

6

—

—

1

•2

4

1

—

-3 ■

,,

83

;)

2

4

—

—

5

9

3

4

—

„

84

4

1

3

—

1

4

6

—

2

—

0)

,,

85

19

—

11

—

—

10

11

3

6

—

^"

'08,

101

24

2

16

—

3

14

26

17

12

—

, ,

126

11

—

B

—

4

4

—

—

—

—

,,

127

38

4

17

—

5

6

6

5

3

—

3

,,

128

16

—

8

—

2

11

6

2

1

—

i^

,,

133

68

—

30

—

12

25

48

31

37

1

o

y

,,

134

30

2

12

•2

1

13

11

5

6

—

1 <

,,

135

102

15

51

—

15

49

85

42

49

—

,,

136

58

8

23

1

7

28

46

21

25

1

J,

137

22

4

6

—

3

12

11

5

4

—

tCfl'

,,

139

41

3

22

—

7

19

28

14

10

—

,,

159

37

4

16

—

8

13

44

14

10

1

' 'oy,

4

46

4

17

1

4

25

54

19

14

2

'08

89

101

7

56

—

14

39

45

24

15

1

,,

93

66

9

42

—

8

23

37

11

14

1

,,

114

73

4

51

—

1

23

—

—

—

—

Rep

. /r,

p. 14

1185

86

605

2

114

499

279

115

128

1

Tot

als

1988

158

1018

6

212

843

781

341

354

8

K. C. PUNNETT

101

TABLE VII.

Families heterozyyous for s/erility and axil.
(Nature of matins FD :• fd.)

Light axil

Dark axil

Reference

^

Number

Fertile Sterile

"09, 1

49 2

„ 2

62 1

'10, 62

114 2

•11, 67

69 3

'12, 102

24 —

„ 104

32 —

„ 108

7 —

„ 109

14 —

Fertile

SterUe

2

11*

1

18

3

30*

3

24

—

5*

—

11*

—

1*

—

7*

Totals 371 8

From Report IV, p. 16 627 27

9
17

107
214

Totals

E.Tpectatioii on 15 ; 1 basis

*These six families ...
Exprctation 15 ; 1 basis

The rest

Expectation 15 : 1 basis

998

35

26

321

993

4H

42

303

Dark

axil

Light

axil

ertile

Sterile

FertUe

Sterile

240

4

5 ,

65 1

,?.?6

9:5

.9 -.5

69 i

758

31

21

256 1

767

32

32

235 (

TABLE VIIL
Families heterozygous for axil and sterility.
(Nature of mating Fd

fD.

Reference
Number

Dark

axil

Light axU

Fertile

Sterile

Fertile

Sterile

'08

84

95

45

44

—

95

31

12

21

—

97

37

29

31

—

106

53

17

22

—

107

214

95

102

—

108

43

22

29

—

109

81

50

58

—

110

72

27

39

—

111

110

55

49

—

112

177

73

99

—

'10,

56

27

17

18

—

»)

60

126

59

69

—

'12,

48

49

28

27

—

Totals ... 1115 529 608

* Families also heterozygous for B and L.

102

Reduplication Series in Sweet Peas

TABLE IX.

F., families from F., normal fr.rliha e.x. F^ plants hc/nrozyi/ons for
slerilUy and cretinism.

(Nature of ;<', mating = Nf nF.)

Norma

I'l-etin

Normal

Refer

ence

, '

-^

, '—

— ,

Reference

/ '

-^

Number

P'ertile Sterile

Fertile

sterile

Number

Fertile Sterile

'12,

70

37

22

34

1

'12,

72

X

xi

71

22

9

9

2

,,

75

X

X

77

X

X

x'

~

,,

81

X

X

79

X

X

X

—

,,

97

33

12

SO

8

f'>

1

—

,,

98

X

X

82

12

VI

8

—

86

17

10

0

—

87

49

11

10

1

'12,

74

GC

—

88

11

7

0

—

,,

78

16

—

89

X

X

X

—

,,

83

X

—

90

24

11

13

—

„

84

42

—

92

11

7

5

—

,,

85

30

—

93

X

X

X

—

,,

94

X

—

99

12

7

3

—

95

X

—

■12,

70

90

20

30

34

■12,

73

X

91

20

4

5

3

TABI

.E X.

90

22

F^ families

from,

/*., pai

•ent, '1

2, 7

6 (Table

IX).

Rpfprpiipp

Normal

(.'retir

Number

Fertile ^

iterile

Fertile

sterile

'13

38

16

8

9

12

,,

39

27

17

9

14

,,

40

20

7

4

9

))

41

12

6

6

9

,,

43

20

7

4

1

>'

45

37

9

10

2

,,

46

18

7

7

2

, ,

42

29

—

19

—

)'

44

(al

four classes

present)

,,

47

23

4

5

4

,,

130

0

1

1

1

Fertile Sterile

4 —

X —

14 —

8 —

X —

' The cross x denotes that the type of plant indicated occurred in these families
though the actual numbers were not determined. In all such cases at least 30 individuals
were examined. The sign — denotes that the class of individual under which it is placed
was not found.

R. C. PUNNETT

TABLE XI.

Cretin

Light

Reference

, — -

— ,

.

Number

Fert.

Ster.

Fert.

A. Fo I'x

'13, 48

102

49

31

DfN X dFn

„ 49

20

17

5

50

14

13

5

51

40

27

8

Dark

Light

b'ert.

Ster.

Fert.

ster

27

6

24

2

4

—

10

—

3

1

6

~

10

1

17

—

176 106

49

44

8 57

B. Fo ami

'13,

52

12

1

1

6

7

F,e.T

53

17

2

—

12

6

DfN X DFn

113

25

—

0

9

15

114

31

1

1

6

7

117

22

—

—

4

3

123

19

1

—

8

8

1 —

126

45

46

13

NOTE ON GAMETIC REDUPLICATION IN PISUM.

By CAROLINE PELLEW,
Minor Student of the John Innes Horticultural Institution,.

It was shown by Vilmorin and Bateson' that when a normal
culinary pea having tendrils (T) and round seed {R) is crossed with
the "Acacia" variety in which the tendrils are represented by leaflets
(t) and the seed is wrinkled (r), j)artial coupling between T and R
occurs in the gametes of F^. The numbers suggested that the system
of coupling was 63 : 1. Further investigation of a number of plants of
similar composition derived (in various generations) from this cross
fully confirm the earlier results. From round seeds the numbers
obtained were 1466 tendrilled, 20 acacia, on the expectation 63 : 1
system being 1471 : 15.

From wrinkled seeds the numbers were 15 tendrilled and 564
acacia, the similar expectation being IS : SGI.

Only seeds of which the starch had been microscopically determined^
were used for these results. This precaution is necessary because not
very rarely occasional seeds of each class may on external appearances
be referred to the wrong class. It should be mentioned that the round
and wrinkled seeds, thus determined, though all the offspring of plants
heterozygous for the.se characters, were selected independently of each
other.

In these experiments the crosses were in the form TR x tr, and the
coupling was

6STR -.iTr-.ltR: 63ir.

Subsequently crosses were made in the form Tr x tR, a tendrilled

1 Proc. Roy. Soc. 1911, 84 B, p. 9.

'' By the method introduced by Gregory, R. P., New Phyt. ii. 1903, p. '226.

106 Gametic Reduplication in Pisuni

variety having wrinkled seed being crossed with a round-seeded Acacia.
The object of this cross was to see whether among the gametes of F^
repulsion between T and R would occur'. The F„ seeds were sorted
into round and wrinkled by microscopical examination, and the result
showed that repulsion occurred. The round seed produced 502 tendrilled,
270 acacia. The wrinkled seed produced 264 jjlants, all tendrilled.

The repulsion is presumably partial ; but if, as is likely, the gametic
distribution is \TR : 687V : 6StR : 1 tr, only one plant in 16,384 would
be wrinkled and acacia, so that any proof of this prediction is beyond
the scope of practical experiment.

In view of the possibility that factors other than roundness might
couple with the factor for tendrils, crosses have been made in which
various factors have been introduced with the tendrilled and acacia
characters. No signs of coupling or repulsion have been observed in
F.2 from such crosses. Among the pairs of characters so tested were
tallness and dwarfness, yellow and green cotyledons, purple and white
flowers, glaucous and emerald f(.)liage, fasciated and normal growth.

' Bateson and Punnett, Journal of Genetics, i. 1911, p. 293.

PRELIMINARY NOTE ON SOME EXPERIMENTS
WITH A POLYMORPHIC PHASMID.

By J. C. F. FRYER, M.A.,

Fellow of Gonville and Gains Coller/e, Balfour Student in the
University of Cambridge.

Among a large brood of " stick-insects," reared from the egg by
Mr E. E. Green, Government Entomologist in Ceylon, it was noticed
that, though the ^/"s were all similar, two distinct types of ^ were
present. The case seemed to merit a detailed investigation, the
preliminary results of which are presented in the following notes.

The insect in question is a typical apterous "stick-insect," the
general appearance of which can be seen at once from the plate
(Plate III); the special characters studied in the experiments were
firstly the colour of the adults, and secondly the presence or absence
of pointed conical horns on the head (Figs. 4, 5). With respect to
these characters the ,/ never varies ; it never has horns, is always
dark chocolate brown in colour and is a much more slender insect
than the female. In the original brood two forms of $ were observed,
the one being horned and green in colour, while the other was hornless
and yellow.

This dimorphism in the ? in relation to the presence and absence
of horns leads to a difficulty in determining the specific identity of the
insect. It belongs to the division Clituiunini of Brunner v. Wattenwyl
and Redtenbacher, but falls into the genus Glitumnus Stal. or Ciinicidina
B. V. Watt, according to whether the % is without horns or possesses
these appendages. Both Westwood in his Catalogue of the Phasmidae'
and Brunner v. Wattenwyl and Redtenbacher in their celebrated

' Catalogue of Orthopterous Insects, Phasmidae, p. 9, PI. 5, Fig. 2.

10!^ Some Expcriiurnts with a Pohiworphic Phasmid

monograph' of the group express a distrust of the character, and it
is now evident that the genus Cumculina as described by the second
author must be dropped. The hornless $ has not been satisfactorily
determined, but the horned $ appears identical with Bacillus cuniculus
Westw., and it will be sufficient for this paper if the subject of the
experiments is assigned to Clitumnus sp ?, jirobably Clitiimnvs cuniculus
Westw.

The first two experiments consisted in isolating a horned green ^
and a hornless yellow $ from the original brood and, as the (/s and $s
had been left together after reaching maturity, it is assumed that both
these $ s had paired.

The following table shows the distribution of the two pairs of
characters among the descendants of these two $ s and their progeny.

Horned Hornless Horned Hornless
Brood green green yellow yellow

Number Parentage Males female female female female

1 Horned sreen ? x j uuknown 13 3 3 10 8

2 Horuless yellow ^ x i unknowu 3 — — — 2

3 Hornless preen ? 1 x ,j 1 48 — 38 — 42

5 Homed green Jlx<fl 30 3 8 8 11

6 Horned green ? Parthenogenetio — 12 10 — —

7 Hornless yellow ? 1 x j 1 Died young. Some ? s with horns

8 Hornless yellow ?lxcrl 10 — — 1 7

9 Horned yellow ? 1 x ^J 1 46 4 6 18 23

10 Horned yellow 9 1 x <r 1 24 4 2 17 1

11 Hornless yellow ? 2 x rf 2 47 — 12 — 43

Note. The number following an individual shows the brood from which it was taken.
Brood 4 died young and is omitted. In Brood 10 the two hornless green ? s died when
immature and there is some doubt as to their colour.

These results are puzzling in inanj' wajs, yet certain facts stand out
clearly. It is evident that the characters of the presence and absence
of horns are Mendelian, as also are those of colouration, and with respect
to both it is noteworthy that no intermediates occurred, .so that every
individual could be assigned definitely to one of the four classes. The
connection between the possession of horns and the green colour,
suggested by the original brood, evidently does not exist, and the
two pairs of characters are probably quite independent of each other.
Finally, and most important, is the definite evidence in brood No. 6
of the segregation of Mendelian factors in parthenogenetic reproduction.
There appear to be no previous records of this phenomenon and con-
sequently confirmation is desirable ; at the .same time the absence of

' Die Iiixckteiifamilic der Phiismidcn, ji. IIIG.

J. C. F. Fryer

109

(/s and the fact that the segregating character is structural seem to
be strongly in favour of the correctness of the result.

As a working hypothesis for the explanation of the experiments as
a whole it may be supposed that horns are caused by the presence of
a factor H, in the absence of which, represented by h, the insect is
hornless. Similarly as yellow colouration appears to be dominant over
green it may be due to a factor G, in the absence of which, c, the insect
is green. Then if the sex factors are represented by MM in the J*
and MF in the % it must be supposed that the combination MM
inhibits the appeai'ance of the female secondary sexual characters.

Under this scheme the (/s and $s must have the following con-
stitutional formulae.

Males

Horned yellow
females

Horned green
females

Hornless yellow
females

Hornless green
females

MMHHCC

ilFHHCC

—

—

—

MilHhCG

MFHhCC

—

—

—

MMHHCc

ilFHHCc

—

—

—

MMHhCc

MFHhCc

—

—

—

MMHHcc

—

MFHHcc

—

—

MMHhcc

__

MFHhcc

—

—

MMhhCC

—

—

MFhhCC

—

MMhhCc

—

—

MFhhCc

—

MMhhcc

—

—

—

MFhhcc

Brood 1 arose from a horned green % and contained (a) about equal
numbers of horned and hornless % s, (h) green % s and yellow J s in the
ratio of 1 : 3.

As regards horns the parent J was evidently heterozygous for H
and the parent (/ contained hh, lacking H altogether.

The colour problem is more difficult and can only be explained by
supposing that the $ paired more than once, which was possible, since
it wa.s left for some time in a cage with numerous (/"s. The formula
of this $ is MFHhcc and if it paired first with a (/" of constitution
MMhhCc and then with one MMhhCC an excess of yellow offspring
would be produced.

Brood 3. A hornless green $ from brood 1 paired with a ^ of the
same brood and gave approximately equal numbers of hornless yellow
and green 2 s. The parent $ being MFhhcc, the parent ^ must have
been MMIiliCc and the result is an equal number of yellow ^s MFhhCc
and green $ s MFhhcc.

Brood 5. A horned green ? fi-om brood 1, paired with a J' of
brood 1, gave 11 horned $s to 19 hornless $s, and 11 green ?s to

Juiirn. of Gen. in 8

no Some Experiments ivith a rolijmorphic Phasmid

19 yellow $s, nuniljeis which are difficult to reconcile with the
hypothesis. The most probable cross is MFHhcc x MMhhC'c which
ought to have given equal numbers of each class.

Brood 6. A horned green $ of brood 1 parthenogenetically gave
approximately ccpial numbers of horned and hornless green $ s. The
parent was evidently heterozygous for H and the offspring would
presumably be MFHhcc and MFIiIicc.

Broods 7 and 8 are only of interest as showing that horns can be
introduced by the male parent.

Brood 9. A horned yellow $ of brood 1 by a </• of the same brood
gave 22 horned $ s and 29 hornless, while as regards colour there were
10 green $s to 41 yellow.

The cross which would give the nearest result to this would be a
? MFHhCc by a </ MMIihCc giving equal numbers of horned and
hornless $ s and green $ s to yellow $ s in the ratio 1 : 3.

Brood 10. A horned yellow $ of brood 1 by a </ of brood 1 gave
21 horned $s to 3 hornless, and 18 yellow ?s to 6 green. This is
another doubtful brood, but the most probable cross to have produced
it is MFHhCc x MMHhCc, giving 3 horned to 1 hornless and 3 yellow,
1 green.

Broods 2 and 11. In brood 2 a hornless yellow ? gave an Fi
containing only hornless yellow $s but in brood 11, the F. generation,
both hornless yellow and hornless green $s appeared — 12 green,
■iS yellow. The first cross may have been either MFhhCc x MMhhCC,
or MFhIiCC X MMhhC'c, and in either case the F^ contained individuals
of constitution MFhhCo and MMIihCc, which if paired would give 3
yellow $ s, 1 green % .

As a whole it is evident that the numbers realised in the experi-
ments do not agree well with those demanded by the hypothesis, but
in this connection it is necessary to point out that there was a very
high mortality during the early stages of the insect's life, which might
easily have disturbed the relative numbers, in which the various forms
of % occurred. It is hoped however that further experiments will be
made with the species and the results, especially those to be obtained
from broods produced parthenogenetically, should be of gi'cat interest.

Finally an expression of gratitude must be made to Mr E. E. Green,
who provided the material for experiment and thanks are also due to
Professor Punnett and Mr L. Doncaster for much advice.

JOURNAL OF GENETICS, VOL.111 N? 2,

PLATE 3

E Wilson .■W

University Press, Cambridge

J. C. p. Fryer 111

DESCRIPTION OF PLATE.

Fig-

1.

Clitumnus sp. ? cf

Fig.

2.

„ ,, ? horned j'ellow.

Fig.

3.

,, ,, ? hornless green.

Fig.

4.

Side view of head of hornles.s green ?

Fig.

5.

,, ,, horned yellow ? .

Note. The species is probably Clitumnus cuniculus Westw. The exact shade of green
and yellow in the two forms of ? is not accurate, as the illustrations were made from
dried specimens and were coloured from memory only.

THE SEGREGATION OF FECUNDITY FACTORS
IN DROSOPHILA.

By EDWARD N. WENTWORTH.

The inheritance of qualitative characters according to the Mendelian
scheme is so well established that even the practical breeder questions
no longer, but quantitative differences are still somewhat slow in
yielding evidence of their accord to this system. Nilsson-Ehle, Shull,
East and other plant breeders have indicated the means by which such
inheritance may take jilace, while Phillips has shown an interesting
example in size in duck crosses. The heredity of fertility is a subject
of practical importance in manj' species, and the problem resembles
those which the investigator has to attack in the genetics of size, vigor,
etc. Pearl has made an excellent contribution to the knowledge of
the subject in his recognition of a sex-linked fecundity factor in the
domestic fowl but beyond this paj)er little work has been done.

The inception of the present experiment was rather accidental.
Early in September 1912 mice gained access to the writer's cultures
of Drosophila and either devoured or liberated nearly everything.
From the wreckage three pupae of orange-eyed flies were rescued
which hatched one male and two females. As one female died before
mating the idea of making an experimental study to see what would
come from inbreeding was conceived. The foundation pair were then
average flies in appearance and possessed of the orange-eyed character.

The -F] generation, if it may be so termed, was composed of
126 flies. Fi'om these four lines were started. The largest and most
vigorous males were mated with the largest and most vigorous females,
the rather weak and small flies were mated together and two lines of
medium type, one based on size and the other on apparent vigor
were also established. These divisions were not based on any actual
measurements, but rather on general appearance and impression.
Fortunately the divisions were justified although later generations

114 Sef/regation of Fecundity Factors in Drosophila

showed that tlieir completeness depended on hick and not on any
knowledge possessed by the writer.

Table I presents the data fioni Line I, the strain lacking in vigor.
Four pairs were mated, they being respectively designated as A, B,

TABLE I.

Breeding

Record q

f Line J.

(A

li7ie

lacking vi

gov

and

high

fecu

ndity. )

Pair A

Pair B

Pair

C

Pail

• D

F,

38

flies

1
1

1
1

34

flies

1
t

51 fl

ies

28 flies

1

1

1

1

1
1

1

1

F-s

30

41

28

32

43

39

36

38

1
r-S
1 1

1

s

1
1 1

1

1

1

r-

s

1 1

F4

32

1

26

1

21

1

34

1

32

1

26

t

32

1

35

36

)

32

1

28

1

31

1

26

1

30

1

32

1

29

1

Fs

1
30

1

1
24

1
26

1
25

1
34

1

1
31

1

1
26

30

1

1
42

1

1
26

1

1
28

1

1
36

1
25

1

1
18

1

1
34

1

40

1

F,

1
32

1

30

29

31

1
30

1

1
26

1

1
34

1

1
28

1

1
44

1

1
30

1
34

1

40

1

1
31

1

1
died
out

31

1

1
41

1

F7

26

1

21

1

23

1

24

1

23

1

21

29
1

34

1

39

1

24

1

44

1

39

1

27
1

25

1

32

1

F,

1
24

1
25

1

1
30

1

1
22

1

1
24

[

28

1
28

1

1
24

1
46

1

1
19

1
60

I
42

1

1
24

1
21

1
34

F,,

25

1

1
23

1

1
19

1

26

1

I
21

1

25
1

1
30

1

19

1

1

30

1

26

1
30

1
39

1

21

1
24

1

27

1

Fn,

1
22

1
25

23

1
21

1
19

1
27

1
24

1
26

1
36

22

24

1
34

18

1
20

1
28

G and D. The table shows the low state of their fecundity and also
the degree to which they bred true. Placed on the basis of the
means, for comparative purposes, the last eight generations rank as
follows :

^3

Mean 35-87J

F.

„ 30-1

F,

„ 29-7

F,

„ 32-73

Fr

„ 28-73

F.

„ 29-4

F,

„ 25-7

^.0

„ 24-6

The mean of the entire line, nine generations, is 29-5 and the
standard deviation is 7-31, giving as coefficient of variability 28-17.
The coefficient of variability is of little value as a means of comparing
the different lines owing to the fact that the units of variation are the
same and the means so widely separated. Thei-efore it would seem

Edward N. Wentwortii 11;")

that the standard deviation is the best method of comparison. The
interesting point is the relative constancy with which Line I breeds
true for fecundity.

To show that this low type is absolutely distinct let us briefly
compare Table I and Table II. While the mean for the low line is

TABLE II.

Breeding Record of Line II. {Strongly Vigoroii.i.)

Pair A Pair B Pair C Pair D

F2 116 132 98 141

I -^ -I I -^ 1 I ^ 1 -I -^ n

I II II I I I

F3 140 128 110 131 112 122 152 135

I I I

n ^ ^ h ^

I I I

^4 161 127 134 117 144 136 128 137 120 129 116 129 147 160 118 151

I I I I I I I I I I I I I I I I
Fa 142 138 127 141 116 134 139 151 138 114 101 145 126 154 136 113

II II II II
^'o 154 136 127 114 141 104 145 162

I I I I II II

F: 136 156 112 144 152 94 139 134

I I I I I I I I
f, 163 147 141 152 127 96 136 157

II II I I I I
Fg 157 138 134 133 141 97 132 172

I' I II I I I I

Flo 149 14* 129 156 132 100 141 ISO

29-5 the mean for the high line is 135'86. The means of each genera-
tion when compared as in Line I are as follows :

F.

Mean

128-75

F,

134-62.5

F,

>i

132-1875

F,

,,

135-625

F,

„

133-375

Fs

,,

140-125

F,

„

138-000

F^o

J)

141-375

The mean for the entire line is 135-86 and the standard deviation
18 3. This gives a coefficient of variability of 13-47, lower than the
coefficient for the low line although the range of variation is obviously
higher as evidenced by the standard deviation. Suffice it to say,
however, that the two lines are quite distinct, they overlap in nowise

116 Segregation of Fecunditg Factors in Drosojyhila

and have each segregated from the common pan: No deterioration
appears, as F,„ is higher than any preceding generation.

The individuals from which Table III is prepared were descended
from four pairs apparently medium in size, while Table IV shows the

TABLE III.

BreecHny Record of Linf. III. {Medium in Size)

Pair A Pair B Pair C Pair D

F-i 81 72 79 84 «

rn r^-i r^-i r^

F3 36 137 79 65 112 75 81 87

II I I I I

I I I I II

n n

Fi 31 60 78 142 72 86 34 77 65 121 68 82 87 144 150 138

I I I I I I I I I I I I II II
Fji 24 71 74 lO'J 136 150 26 84 71 86 38 141 28 128 146 127

II I I I I I I
Fo 19 68 138 68 42 54 28 86

II ill! I I

F^ 31 89 161 75 68 20 21 74

II I I I I I I

Fs 26 102 124 56 80 34 24 69

II I I I I I I

Fa 22 86 152 18 144 32 20 77

II 1 I I I I I

/•',„ 25 41 148 24 127 40 23 81

TABLE IV.

Breeding Record of Line IV. {Medium in Vigor.)

Pair A

Pair

B

Pair C

Pair D

F.

75

1

1

80

64

78

1

1

t\

78

r 1

72 76

1 1

1

1

44

r n

1 1

41 35

1 1

g

1
1
12

1

1

87

t

71

1

69

74

85

1

J-\

1

1
71 139

1 1

1

1 I
82 69

1 1

J
30

1
67

74 77

1 1

1
1
68 70

1 1

1
1

78 82

1 1

Ft,

1 1
68 25

1

78 64

1

1
28

1

1
72

1
40

1
71

70

1

29

112 56

71 119

1 1
1.34 70

1

Fo

1
29

1

1
72

1

1
32

02

1

1
112

135

74

1

1
71

1

F-;

1
21

1

1
141

1

34

1
08

1

101

120

1

1
05

I

1
20

1

F,

1
26

1

1
134

29

1

1
51

1
119

1
124

1

1

81
1

1

55
1

F,

1
19

118

1

1
25

1

34

126

1

1
74

1

77
1

1

82
1

Fm

22

1
137

1
27

25

1
117

76

1
74

1
73

Edward N. Wentworth 117

descendants of four pairs medium in apparent vigor. Since both show
the same qualitative results they will be combined for the comparison
with Tables I and II. The means of the separate generations follow :

i^. Mean 78-875

F,

., 81-1875

F,

„ 83-6875

F, ..

„ 68-125

F,

„ 60-0625

F, ,

„ 71-25

F,

„ fi9-125

F^o

„ 66-25

The mean of the entire lot is 72 06, and the standard deviation
is 37-1618. This gives the largest coefficient of all, 51-27, and from
the standpoint of the coefficient would offer evidence of a 1 : 2 : 1 ratio.
That this is merely apparent will be shown a little later in the paper.
Summarized we get the following table.

standard Coefficient

Mean Deviation of Variation

Low line ... 29-5 7-3 28-17

Medium line ... 72-06 37-2 51-57

High line ... 135 -80 18-3 13-47

A segregation of fecundity factors is clearly evidenced and the
supposed weaknesses from inbreeding are shown up in their true light
as the mere segregation of factors for lower vigor.

In order to confirm the idea of the relative separateness of these
fecundity factors a reciprocal cross was arranged between flies from
the hatch of 142 in Line III, pair A, Ft, and flies from the hatch of
30 in Line IV, pair C, F^. Table V shows the detailed results of this
work, the confirmation being absolute. The interesting point brought
out is the fact that the male, whether he come from high or low lines,
apparently in nowise influences the eggs laid by the female with whom
he pairs, though marked differences, apparently due to segregation, may
occur among his female descendants.

Further evidence on the nature of such segregation was obtained
by breeding all the pairs available in one hatch from an apparently
intermediate pair. The pair selected was from Line IV, pair B, F^,
Hatch 82. Thirty-nine males and forty-three females appeared and

lis iSef/regatioH of Fecundt'tf/ Factors In Drosoj>/n'/a

TABLE V.

Data from Reciprocal Cro^acs of Strains IVnih and Loio in Fecundity.

Ft, i From Line III, Pair A (liatoli 142), ? From Line IV, Pair 0 (liatL-li ;^0)

I I

I 1 I

H4 flies (14 J s and 20 ? s)

Pair T Pair I] Pair V Pair .\ Pair 1' Pair Z

32 70 84 79 35

' An r-^n r^ r^n

II II II

I

n n

Ft 28 34 36 86 121 75 141 78 27 64 82 78

II II II II II II

J's 24 29 27 116 126 81 124 91 23 76 79 80

Reciprocal .

Fr, 1 From Line IV, Pair C (hatch 30), i From Line III, Pair A (hatch 142)

' J

129 flies

(61 J

s and 07 ? s]

1

Pair T

Pair (7

Pair r

Pair X

Pair Y

Pair Z

F,,

86

31

134

69

80

T2

i 1
79 134

r

1

31 78

1

1 1

1 1

F,

26 39

131 142

76 81

116 81

Fh

1 1
80 121

32 29

85 137

1 1
34 84

1 1
122 86

131 42

from this thirty-nine cultures were started. The progeny of each of
the pairs is listed in Table VI. Instead of three simple groups which

TABLE VI.

7'A« Record of all the Offspring from one Pair of Flics from
Line IV, Pair B, F„ Hatch 82.

Pair 12 3 4 5 6 7 8 9 10 11 12 13 14 15

Number 53 19 99 131 74 38 142 76 40 102 57 129 122 59 42

Pair 16 17 18 19- 20 21 22 23 24 25 26 27 28 29 30

Number 77 108 112 78 90 60 88 45 87 107 80 62 87 104 64

Pair 31 32 33 34 35 36 37 38 39

Number 81 101 81 65 96 84 103 85 82

might be expected from the previous calculations there are at least

Edward N. Wentworth

lUt

seven. In fact the interpretation on a three-factor basis corresponds
very well to the distribution of these pairs. Assuming twenty to be
the niininiuni fecundit}', since it is approximately the lowest that
segregated out, and one hundred and forty the upper limit with rather
wide fluctuations, in all groups the intervening number divides readily
into six parts of twenty each. An hypothesis involving three pairs
would permit of six factors in the individual homozygous for high
fecundity, these factors to be of twenty each. If this were the case,
then the expectation and actual results would be as follows, the actual
pairs being somewhat arbitrarily distributed owing to their lying in-
termediate between the groups of twenty.

Number Factors

Six

Five

Four

Tliree

Two

One

None

Fecundity Number, Offspring

140

120

100

80

00

40

20

Expected Number

0-6

3-G

9-0

12-0

9-0

3-6

0-6

Actual Number

1

3

9

14

7

4

1

Number produced by each group on

142

131

99

74

53

38

19

the basis of which the above calcu-

—

129

102

70

57

40

—

lation was made (Cf. Table VI) . . .

—

122

108

77

59

42

—

—

—

112

78

60

45

—

—

—

107

80

02

—

—

—

—

104

81

04

—

—

—

—

103

81

05

—

—

—

—

90

82

—

—

—

—

101

84
85

—

—

—

—

—

87
87

—

—

—

_ _ _ 90 _ — —

Circumstances prevented the breeding out of these different groups,
so no definite confirmation of the above hypothesis can be offered.
This does not account for the many progenies that numbered above
140. It is possible that at least four of these factors are concerned
in the race instead of three, but the particular pair under consideration
apparently bore only three of theni. The data used in Tables I^IV
inclusive more nearly fit a four-factor hypothesis than the three-factor
reported for Table VI. No evidence of sex linkage of fecundity factors
appeared.

120 Segregation of Fecundltg Factors in DrosojyJtila

LITERATURE CITED.

1908. East, E. M. " Inbreeding in Corn." Conn. Agrl. Exp. Sta. Report, VMl—

1908, pp. 419— 4 iS.
1910. . "An Interpretation of Variation that is Apparently Continuous.'

Amer. Nat. Vol. .XLiv. pp. 739 — 746.
1912. and Haye.s, H. K. "Heterozygous in Evolution and Plant Breeding."

B%d. 243, Bureau of Plant Industry., U.S. Dept. of Ayrkultnrc, pp. 58.

1909. Nilsson-Ehle, H. " Kreuzung.suntersucluuigen an Hafer und Weizen."

Lunds Universitets Arsskrift, Vol. v. No. 2, pp. 122.

1912. Pearl, Raymond. " The Mode of Inheritance of Fecundity in the Domestic
Fowl." Journal E.vperimental Zoology, Vol. xill. No. 2, pp. 153 — 268.

1912. Phillips, J. C. "Size Inheritance in Ducks." Journal Experimental
Zoology, Vol. XII. No. 3, pp. 369—380.

1908. Shdll, G. H. "The Composition of a Field of Maize." American Breeders'
Association Reports, Vol. iv. pp. 296 — 301.

MEN DELI AN SEX-FACTORS IN MAN.
By J. A. JENKINS.

That the numerical equality of the sexes in so many types may be
most simply explained on the assumption that one sex is homozygous,
and the other sex heterozygous for sex-factors ("maleness" and "female-
ness"), is I believe commonly granted. Again, several instances are
known in which there appears to be (during gametogenesis) "repulsion"
between a sex-factor and a character-unit, resulting in a " sex-limited "
descent of that character ; but no clear case of " coupling " ia similar
circumstances has yet been brought forward. A necessary condition for
either repulsion or coupling to take place is the heterozygosis of both
factors in the parent concerned (Bateson, Mendel's Principles of
Heredity, C. U. Press, p. 1.51). In sex-limited cases the following
general considerations arise :

(1) As by our hypothesis one sex is homozygous and the other
heterozygous, the dissimilar sex-factor borne by the latter parent must
in all cases be dominant : also, it is in the gametogenesis of this parent
that we look for the " sex-limitation."

(2) The process in such cases being due to repulsion existing
between dominant factors, we expect to find among the progeny that
individuals heterozygous in sex bear the recessive character (of another
pair of factors) more frequently than individuals homozygous in sex.

Applying these rules to human colour-blindness, and taking colour-
blindness, n, to be recessive to normal vision, N (as shown to be the
case by Doncaster, Journal of Genetics, Vol. i. p. 378), we see that
probably in Man, the female is homozygous and the male heterozygous

122 Mendeliau Sex-Factors in Man

for sex-factors, and that " vnalcness " is dominant to " femaleness."
Individuals (zygotes) may therefore be represented as:

Normal female NN i ?

Colour-blind female nn^ t

Heterozygous (apparently normal) female Nn ? ?

Colour-blind male nn cf ?
" Normal " male n i . N ■}
(repulsion existing between N and i )

The various equations become :

(a) Normal female x CB male.

(b) Heterozygous female x " normal " (really heterozygous) male.
Nn^ ? x/.J".iV? =i>»^^ +Nn^^ + i\"« ? ? +NN^ ?.

(c) Heterozygous female x GB male.

Nni^ X nnj' ? = >m,f ? + NnJ' ? + nn $ $ -f iV/( $ ? .

(d) CB female x " normal " (really heterozygous) male.

H«? ? xn^.N^ =nn^% +Nn%%.

If the above view of the sex-composition of Man be accepted, it of
course follows that the sex of offspring is determined, not by the mother,
but by the father, in Man.

{Received 1 Jan. 1913.)

ON THE RECOGNITION OF THE INDIVIDUAL
BY HAEMOLYTIC METHODS.

By CHARLES TODD, M.D.

The discovery and elaboration of the so-called "Immunity Reactions"
show signs of proving one of the greatest advances made by biological
science during recent years, and though discovered originally by
bacteriologists in connection with the study of disease, and for a time
practically limited to bacteriological research, the significance of these
reactions has since overflowed into other fields, and they now constitute
our most valuable method for the identification of the proteins.

For the benefit of those unacquainted with the subject of immunity
we may briefly instance one or two concrete examples indicating the
nature of some of these reactions. If, for example, a rabbit is injected
with the white of a hen's egg, it is found that after a certain interval
of time the blood-serum of the rabbit which has been so treated has
acquired the jjroperty of causing a precipitate when added to a solution
of hen's egg albumen. This reaction is extraordinarily sensitive — a
dilution of even 1 in 100,000 of the albumen still giving a precipitate.
It is, moreover, specific, that is to say, the serum, if used in suitable
dilutions, only gives a precipitate with the albumen of the hen's egg
and not with that of other birds such as ducks, geese etc.

In the same way if a rabbit is injected with human blood, the
serum of the rabbit acquires the property of causing a precipitate
in solutions of human blood and this constitutes a valuable method of
distinguishing human from other blood — a method now in regular use
for medico-legal purposes.

The statement that these reactions are specific requires qualification
as this is only the case in certain dilutions. If strong solutions are
used the serum causes precipitates, not only in solutions of the blood

124 Recognition of the Individual by Haemohjtic Methods

of the animal used for the injection, but also in those of closely allied
species. As the degree of dilution is increased, however, the pre-
cipitates given with allied species gradually diminish until a point is
reached where the serum only gives a precipitate with the hlo(jd of
the species for which it has been prepared. Taking advantage of this
fact, the method has been used to investigate the relationship of the
various animal species to one another and most interesting results in
this direction have been obtained, notably by Xuttall and Uhlenhuth.
These observations have shown the relationship of man to the ape, and
the fact that the apes of the old world are more closely akin to man
than those of the new.

These methods are extraordinarily delicate and Uhlenhuth has
shown that by their help it is quite possible to distinguish between
such closely allied bodies as the albumen of the fowl's egg and the
albumen of the blood of the same bird — a difference which is, of course,
far beyond the reach of our present chemical tests.

It is impossible here to go into the many interesting results which
have been obtained by means of the pi-ecipitin reaction, but reference
may be made to the work of Uhlenhuth and Roemer on the protein
substances of the crystalline lens of the eye. These observers found
that the crystalline lens of all animals, right down the zoological scale
as far as the fishes, possesses a biologically similar protein which more-
over appears to have no connection with the blood proteins of the same
animal — a serum prepared for instance with the lens material of the
ox gave a precipitate equally with solutions of the lenses of the pig,
man, fowl, frog etc., but gave no precipitate with the blood of these
animals, so that the lens must therefore be regarded as regulai-ly
consisting of a protein foreign to the organism.

Another well known inimuiiity reaction which is now in everyday
use for the practical diagnosis of typhoid and Malta fevers etc. is the
so-called "agglutination test" which depends upon the fact that when
certain bacteria (e.g. the typhoid bacillus) are injected into an animal
the serum of the animal so ti'eated becomes agglutinating for the
species of bacillus injected, that is to say when added to a suspension
of the bacilli, it causes these to clump together and fall to the bottom
of the vessel in which they are contained. The exact nature of this
phenomenon is not known, though various explanations have been
suggested; it would appear however that the agglutination is most
probably due to some change in the surface tension existing between
the bacilli and the Huid in which they are suspended.

Charles Todd 125

All the immunity reactions which are known appear to depend
upon the fundamental fact that when a foreign protein is injected
into an animal, the organism responds by the formation of an
"antibody" which is specific for the protein injected. This specificity
is extraordinarily complete and the relation between the protein and
the corresponding antibody has been compared to that existing between
a complicated lock and its key. It appears to be a sine qua non that
the injected substance should be a colloid, and carbohydrates, fats etc.
do not give rise to the formation of antibodies.

If half a dozen different foreign proteins be injected into an animal
at the same time, the corresponding six antibodies duly appear in the
serum ; and it is difficult to suppress a feeling of astonishment at the
perfection of a mechanism which enables the animal organism, within
the space of a few days and with no apparent exertion, to synthesise a
series of new compounds of such extraoi'dinary complexity.

The number of antibodies which are known is now very large
and is constantly increasing, but the above examples are probably
suflScient to give a general idea of the conditions under which
immunity reactions take place, and we may proceed at once to
consider the question of haemolysis with which we are more im-
mediately concerned.

The study of haemolysis, thanks to the labours of Bordet, Ehrlich
and Morgenroth, and a host of other workers, has added greatly to our
knowledge of the phenomena of immunity, and haemolytic methods
now find their place in the routine work of most bacteriological
laboratories. The subject is unfortunately of very great complexity,
and its somewhat appalling nomenclature has hitherto not tended to
render it attractive to workers in other branches. The phenomena in
themselves are however simple, and by keeping to concrete cases and
avoiding as far as possible technical terms, are easily described.

Supposing, for instance, a rabbit is injected with the red blood
corpuscles of the ox, it is found that the serum of the rabbit, which
before treatment had no action on ox corpuscles, acquires after a few
days the power of di.ssolving these corpuscles very rapidly, owing to
the formation of a haemolytic antibody or " haemolysin " in the serum.
We can show that the haemolytic action is actually due to the presence
of this haemolysin, by mixing some of this haemolytic serum and fresh
ox corpuscles and keeping the mixture at 0° C. overnight in the ice
safe and then centrifuging. At this temperature no haemolysis takes
place but the haemolysin is bound by the corpuscles, and is entirely

Journ. of Gen. in 9

126 Becof/nifion of the Imlivkhial hji Haemoh/h'c Methods

removed from the serum, which is now found to be without any trace
of solvent action on fresh corpuscles.

The serum of the rabbit which has been injected with ox corpuscles
resembles the precipitin sera referred to above in that it is not strictly
specific, but in addition to haemolysins for the ox, also contains
haemolysins for animals of species allied to the ox, such as the sheep,
goat etc.

If it is desired to remove these secondary haemolysins, this is easily
accomplished by the method of " exhaustion " or " elective absorption."
Supposing, for instance, we wish to remove the haemolysin for the
sheep, it is only necessary to " exhaust " the serum with sheep's
corpuscles, i.e. to leave it in contact with sheep's corpuscles at 0° C.
and then remove these corpuscles by centrifuging. The serum is then
found to be quite without action on sheep's corpuscles, whilst it remains
still active for the corpuscles of the ox. In this way sera, which are
originally haemolytic for two or three species, can be artificially
rendered haemolytic for only one of them, that is to say they can be
made more specific.

Bordet having shown that the injection of blood corpuscles into
an animal of a different species gave rise to a haemolysin, Ehrlich and
Morgenroth investigated the results of injectiiig animals with the
corpuscles of other animals of the same species and found that in
this case also as a rule a haemoly.sin is formed. Thej' injected a
goat with blood obtained from another individual of the same species
and found that the serum of an animal so treated became haemolytic
for the corpuscles of the individual whose blood was injected, and also
for those of other goats but never for its own corpuscles. (To use their
nomenclature : it became isolytic but not autolytic.)

By injecting goats with goat's corpuscles, thirteen of these isolytic
sera were prepared and a careful study of their properties, by methods
too complicated to be gone into here, revealed the fact that they
all differed fi-om one another, i.e. that they represented different
isolysins.

The investigation of the isolysins was continued by Todd and
White"'' '='■ '^' who had at their disposal 106 cattle which in the course
of their immunisation with cattle plague had been injected with large
quantities of the blood of other cattle.

An examination of the sera of these animals showed that
76 of them were very highly haemolytic for the corjDuscles of normal
cattle and a detailed investigation gave most interesting results.

Charles Todd 127

When one of the sera, was tested on a series of corpuscles of
different normal cattle, it was found that the haemolytic power
of the serum was quantitatively very different for the corpuscles of
different individuals; being very highly haemolytic for some, less so
for others, and again comparatively slightly so for others. In a similar
test made on the corpuscles of the same individuals but with a second
serum the same variations were observed but they did not coincide
with those obtained with the first serum, that is to say corpuscles
which were highly haemolysed by the first serum were often com-
paratively slightly affected by the second and vice versa. When the
tests were extended to other sera similar results were obtained — in
short no two sera ajjpeared to be absolutely alike.

It was next found that if one of these isolytic .sera is exhausted
with the corpuscles of a particular individual ox (A), it remains
haemolytic for the corpuscles of many other individuals, but loses its
haemolytic power for the corpuscles of some other individuals as well
as for those of (.4 ).

If now a second isolytic serum is exhausted with the corpuscles
of the same individual (A) and then tested, its action on the various
corpuscles is not exactly jjarallel to that of the first serum and often
shows marked differences. This result is to be expected as it was
shown by Ehrlich and Morgenroth that two goats each injected with
similar doses of the same goat's blood at the same time gave quite
different isolysins.

As a matter of fact the isolysins formed depend upon two distinct
factors :

(a) The individuality of the injected corpuscles.

(b) The individuality of the animal into which they are injected.

When we consider the enormous number of variations possible in
each of these factors, we see the almost unlimited possibilities in the
resulting sei-a.

In view of the above it should be possible by taking a mixture
of a sufficiently large number of immune sera and exhausting this
with the corpuscles of one individual, to obtain a serum which has no
haemolytic action on these corpuscles but haemolyses those of all other
individuals of the same species and might therefore be made use of for
the identification of this particular individual.

To test this a mixture was made of between 60 and 70 of these
isolytic sera. This mixture was then exhausted with the corpuscles

9—2

128 Kecognitwn of the Individual hy Haemolntic Methods

of a normal ox and then tested on the corpuscles of 110 different
individual cattle. It was found that the mixture was powerfully
haemolytic for the corpuscles of every one of these 110 individuals
but absolutely without action ou the corpuscles of the individual
for which it had been exhausted. A number of other tests was made
by exhausting the serum with the corpuscles of various individuals
and similar results were always obtained, except in the case of close
blood-relations where certain exceptions occur which will be referred
to later.

We may therefore conclude that the red blood corpuscles of any
individual (excluding for the moment the question of close blood-
relations) possess characters which differentiate them quite distinctly
from the red blood corpuscles of any other individual even of the same
species.

Being now in possession of a method of identifying the corpuscles
of a certain individual even in the presence of corpuscles of other
individuals, it was possible to study the question of what happens when
the corpuscles of one individual are introduced into the circulation of
another individual of the same species. Eight oxen were injected
intravenously with the fresh blood of other oxen, quantities of from
two to four litres of blood being injected. In all these animals a
similar course of events was observed, the number of the foreign
corpuscles in the circulation gradually diminishing until they dis-
appeared after a lapse of from four to seven days after the injection.
Shortly after the disappearance of the foreign corpuscles from the
circulation, the blood .serum began to acijuire haemolytic properties.
These experiments emphasise in a most striking manner the definite
individuality of the red blood corpuscles, and we see that the injected
corpuscles are not merely not accepted by their new host, but are
regarded as definitely foreign and in f;ict give rise to the formation
of corresponding antibodies in accordance with the general laws of
immunity.

Having found that it was po.ssible to distinguish the corpuscles
of non-related individuals of the same species several tests were made
on the corpuscles of closely related individuals. A mixture of a large
number of haemolytic sera was exhausted separately with the corpuscles
of a cow and her calf It was found that while exhaustion of the serum
with the corpuscles of the calf removed the haemolysin for the calf
only, exhaustion of the serum with the corpuscles of the cow removed
the haemolysin not only for the cow but also for the calf. A similar

Charles Todd 129

examination of a family of sheep (consisting of the father, mother and
three lambs) showed the interesting ftxct that the corpuscles of one
lamb resembled almost exactly those of the mother, whilst the corpuscles
of the other two lambs had the characters of the father.

From what has preceded we see that it can be definitely proved
by tests made in vitro that in the case of a warm-blooded animal
the red blood corpuscles of any one individual are different from those
of any other individual of the same species (always excepting the case
of close blood-relations). Moreover the corpuscles of one individual
when introduced into the blood-stream of another individual are re-
garded by their new host as foreign bodies and are treated as such.
A consideration of this very striking fact at once leads to speculation
as to whether this is not merely one example of a general law holding
for all the cells of the body. Unfortunately the other cells present
much greater experimental difficulties than the red blood cell, whose
delicate stroma renders it accessible to the quite extraordinarily sensitive
methods of haemolysis ; we have, however, a considerable aniount of
indirect evidence in favour of this view, derived mainly from the results
of transplantation of the various tissues in man and other warm-blooded
animals. Schiine, in his most interesting work {Die Jieteroplastische
•and homooplastische Transplantation), summarises the results of other
observers and also gives an account of his own extensive researches on
this subject. Perhaps the most striking results given are in connection
with skin grafting. It is now generally recognised that whereas skin
grafts replanted on to the individual from which they were taken
(autoplastic transplantation) succeed with ease ; those planted on to
a different individual of the same species (homoioplastic transplantation)
are exceedingly rarely, if ever, successful, unless the two individuals
happen to be blood-relations. Schone woi-king with mice found that
homoioplastic transplantation failed almost as regularly as autoplastic
transplantation succeeded. His experiments with blood-relations showed
that transplantation between brothers and sisters and from the child to
the mother were possible ; but, curiously enough, in the few cases in
which it was tried, transplantation from the mother to the child did
not succeed. This result is particularly interesting in connection with
the haemolytic experiments recorded above on the difference between
the red blood corpuscles of the cow and her calf In the transplantation
of bone Axhausen's most careful experiments show that autoplastic
transplantation succeeds much better than homoioplastic. Rehn has
shown that the same holds good for the transplantation of fatty tissue.

130 Recognition of the Individual hji Haemolytic Methods

and other observers have proved the same fact for the transplantation
of the thyroid gland and the ovary. It is a matter of common know-
ledge that the blood cells, both red and white, live considerably longer
in their own serum than in that of another individual of the same
species and a considerable amount of similar indirect evidence is
available.

In view of the above facts we are probabl3' justified in assuming
as a working hypothesis that, at any rate in the case of the warm-
blooded animals, all the cells of the body of an individual are, so to
speak, stamped with his individuality and are different from the
same cells of any other individual (not a blood-relation), even of the
same species.

REFERENCES.

(1) Todd, C. and White, R. G. rrocecdingx of tJi>: lluytil Soci'cU/, 11, Vol. Lx.x.xil.

1910, p. 416.

(2) . Proceedings of the Royal Society, B., Vol. Lx.\.\iv. liJll, p. 255.

(3) . Journal of Hygiene, Vol. x. 1910, p. 185.

SOME POINTS OF GENETIC INTEREST IN
REGENERATION OF THE TESTIS AFTER
EXPERIMENTAL ORCHECTOMY IN BIRDS.

By C. J. BOND.

In 1906 I published some observations on the result of unilateral
oophorectoni}' in Rabbits (see " Inquiry into some points in uterine
and ovarian physiology and pathology in Rabbits," B. M. J., July 21st,
1906). In this communication it was shown, among other results, that
the removal of one ovary in the female Rabbit is followed by a com-
pensatory overgrowth in the remaining ovary, and further that this
hypertrophy affects both the Graafian follicles or ova-bearing cells and
the internal secretion-forming cells.

At this point it seemed desirable to ascertain whether the same
process occurs in the remaining testis after unilateral orchectomy.

This was found to be the case in Birds but during this experimental
investigation' some further facts were elucidated which seem to me to
have some important genetic interest.

In the first place it was found in both male Fowls and Pigeons
that when one or both testes were removed intracapsularlj-, that is to
say when the testicular substance had been apparently wholly removed
and the capsule alone left, a regeneration of the secreting tissue of the
testis and the tubuli seminiferi took place within the capsule so
evacuated.

In a certain period of time, varying from six weeks to six months,
the whole substance of the normal-sized testis may be thus restored.
Fig. 1 shows the naked-eye view of two testes so regenerated in each
of two cockerels, in one case (22 c.) after six months and in another
(20 c.) after seven months. PL IV, fig. 2 shows a transverse section of

' The actual operations connected with this investigation were kindly performed for
me by Sir V. Horsley, F.R.S., at University College, London.

132 Uxpei'imental Orchectomy in Birds

one of these testes so regenerated under a low magnification, S" obj.
PL IV, fig. 3 shows a portion of the same section under a higher magni-
fication, \" obj. In some sections the regenerated tubuli seminiferi are
seen to be filled with spermatozoa as in the normal gland.

(20 c.)

(22 c.)

Fig. 1. Eegeneration of the Testes after Subcapsular
Castration in Fowls. (From Photo.)

Thus it is clear that a real regeneration of .sperm-bearing testicular
tissue occurs after subcapsular orchectomy and this regeneration must
take place either from the capsule, or more probably from microscopic
fragments of secreting ti.ssue which are left adhering to the capsule at
the time of the operation.

Such evidence as exists goes to show that regeneration of testicular
substance within the capsule after removal does not occur, at any rate
to any appreciable extent in the case of some mammals. This is a
matter of some importance. The domestic fowl has undergone a
process of artificial selection in regard to the egg-laying capacity of
the female sex gland, and probably also in regard to the sjjerm-forming
capacity of the male sex gland, hence it is of interest to find that the
testis of the domestic fowl has great powers of structural and functional
regeneration after partial removal, greater apparently than exists in
mammals.

JOURNAL OF GENETICS, VOL. IN. NO. 2

PLATE IV

Fig. 2. Transverse Section of Testis of Fowl after Eegeueratiou. '6" obj.

Fig. 3. Transverse Section of liegenerated Testis of Fowl, i" obj.

JOURNAL OF GENETICS, VOL. IIL NO. 2

PLATE V

Fig. 4. Cook, Wyauilotte Type, and Hen, Brown Leghorn
Type. Father and Mother of Cockerel 7.

Fig. 5 A. Coclierel 7, and daughter
after Castration.

^l?f="f:ri-

'"*■ - ■ ?;.

Fig. 5. Cockerel 7, and Sister Hen (mated) and
daughter before Castration.

Fig. 10. Shows regenerated ovary in Hen, 1 year
and 9 months after destruction by Cautery.

X Regenerated ovary.

C. J. Bond 133

Having ascertained that reformation of gamete-bearing tissue does
occur after removal in the male Fowl it became a matter of genetic
interest to ascertain whether the gametes which are formed in this
regenerated tissue resemble, in their hereditary characters, the gametes
which are formed by the original gland before removal. I will now
describe some experiments on Fowls and Pigeons which bear on this
point.

Experiment 1.

A Hybrid Silver Wyandotte and Brown Leghorn Cockerel (Cockerel
number 7) was bred of the following Genetic composition :

Brown Leyhoru x Silver Wyandotte

r 1

15 S 16 V

All resemble Wyandotte Mother All resemble Brown Leghorn Father

I I

Cock, Wyandotte Type x 3 Pullets, Brown Leghorn Type

PL V, tig. i.

I

-n

( 8 Wyandotte Type 7 Wyandotte Type

/ 7 Brown Leghorn Type 7 Brow'n Legliorn Type)

PI. V, fig. 5.

Cockerel 7x3 Pullets
Brown Leghorn Type I Brown Leghorn Type

r ]

Is 6?

All Brown Leghorn Type All Brown Leghorn Type

PI. V, tig. o («).

Cockerel 7 x Same Hens

Subcapsular Orchectomy I
Oct. 1908

I —

I I

All Brown Leghorn Type

Thus Cockerel 7 produced chickens of the same Brown Leghorn
Type before and after regeneration of the testis, when mated with
the same hens.

Fig. 6 (p. 134) shows a photo of the regenerated testes, life size, when
the bird was killed in June 1910.

134 Exjwrimental Orchectomi/ in Birds

Fig. 7 shows the iionnal testes of a cockerel aged two years, life size,
for comparison.

Fig. 6. Photo, Life Size. Kegenerated Testes 1 year and 9 luontlis after
Subcapsular Castration. Cockerel 7.

Fig. 7. Photo, Life Size. Normal Testes, Cock, aged 2 years.

C. J. Bond 135

Regeneration of Testes in Pigeons.

Experiment 2.

A Blue Checqiier Fantail was bred of the following Genetic

composition :

<? ?

Black Fautail x White Fantail

I

. ,

J ?

Black and White Fantail x Black and White Fantail

I

r T- ^ -T n

II II

Black Blue Checquer Black and White White

s ?

Blue Checquer x Blue Checquer

r ]

Dec. 1906 to July 1908

12 Blue Checquers

Blue Checquer x Same Heu

Subcapsular Orcheetoniy
Oct. 1908

I

"1

Dec. 1908 to July 1910
17 Blue Checquers (4 with feathered tarsi and toes)

The offspring of this Blue Checquer Cock after regeneration of
the testes are identical in plumage with offspring hatched before
subcapsular orchectomy.

One point should be noted. Among the young hatched before
castration none showed any feathers on the toes, whereas out of the

17 hatched after regeneration of the testes four showed signs of
feathered tarsi.

Fig. 8 (1) (p. 136) shows the regenerated testes (life size) of the Blue
Checquer Cock one year and seven months after castration.

Fig. 8 (2) shows photo of normal testes (life size) of a White Fantail
for comparison.

Fig. 9 (1) shows photo of the testes (atrophied), life size, removed
from a White Fantail aged nine years which had been sterile

18 month.s.

Fig. 9 (2) shows the normal testes of a Fantail three years old of
the same size.

136

Experimental Orclieetomy in Birds

(1)

(-')

Fig. 8. (1) Sliows Regenerated Testes. Life
size of Blue Checquer Cock.

(2) Shows Normal Testes of Fantail. Same
size and age for comparison.

(1)

(2)

Fig. 9. (1) Shows I'hoto Life size of Atrophied
Testes removed from Fantail Cock, age 9 years,
Sterile 18 months.

(2) Shows Normal Testes of Fantail, same
size 3 years old.

C. J. Bond

137

Experiment 3.

A Hybrid Tumbler Fantail Pigeon was bred of the following Genetic
composition :

c? White Fantail x ? Black Tumbler

I

Black and White
do. i

Brown, Blue and White
do. ?

I
Brown

Black Black and White x Brown, Blue and White White
do. i I do. ?

Ill I I II

Blue Almond Blue and Almond and Black and Brown, Blue White

White White White and White

I I

Blue and White x Ahnond and ]Vhite

s I ?

, -L -r

6 Blue and White

.3 Blues

Blue and White
i
Subcapsular Orchectomy,
Sept. 17, 1908

I

1 White (N.B. 2 died soon after
hatching)

Same Hen

I \ 1 I I

III I I

Blue 11 Blue and White 9 Almond 2 Almond and White .5 White 7

N.B. Three died soon after hatching.

From the above table we see that the Blue and White Cock mated
with an Almond and White sister produced, before castration, 10 young
pigeons of which number one only was white. After castration and
regeneration of the testes the same pair produced 34 young pigeons of
which seven were white.

It is interesting to notice that in the smaller number of young
hatched before castration, the proportion of Recessive White to the
rest was 1 in 10, while after castration and regeneration of the testes
the proportion of Recessive White to the rest (in the larger numbers)
was nearly 1 in 5.

Although in the case of the fowl the male contribution to the
zygotes hatched before and after regeneration of the testes were alike
in character, as judged by the appearance of the chickens, this result
may have been due to the fact that only one kind of gametes was being
formed by the testes of the particular bird operated on, and so the

138 Experimental Orchectointj hi Birds

renewed growth of the testes after removal in this bird only led to the
reproduction of the original kind of gametes.

Conclusions.

Such arc the facts brought out by this investigation. Such as they
are, they seem to indicate that the cell divisions of the mother sperm-
cells which provide the new spermatozoa formed during regeneration
of the testis do not take place in exactly the same order, or are not
exactly of the same kind, as those which form the sperm-cells before
removal of the sex gland.

Thus in Experiment 2 among the 17 young pigeons hatched after
castration of the male parent, four showed feathers on the tarsi and
toes, while among the 12 hatched before castration none were so
marked.

Again in Experiment 3 (as has already been stated) the proportion
of Recessive Whites rose from 1 in 10 before, to 1 in 5 after castration
of the male parent.

It is true that in Experiment 1 chickens hatched from eggs fertilijied
by Cockerel 7 when mated with three hens (sisters to cockerel) resemble
the chickens hatched from eggs by the same hens fertilized after the
testes had been regenerated. But the question of the genetic composi-
tion of this bird must be considered. If the bird was homozygous in
respect of this character of colour, that is to say if only one kind of
gametes were being produced before castration, then there is no reason
to think that the increased production of spermatozoa set going, during
regeneration of the testis after subcapsular orchectomy, would result in
the production of gametes of a different kind.

The suggestion is, that in a sex gland where gametes of different
factorial composition are being produced the temporarily arrested and
the subsequently increased cell division due to the stimulus of removal
of the organ would be likely to result in a different rate, or order of
production of, the different kinds of gametes which are being formed
in that particular male organ. This can of course only be judged of by
a comparison of the zygotes which are hatched from eggs fertilized
by these male gametes before and after this stimulus of removal
has been applied.

For this reason it seems desirable to simplify the conditions of the
experiment and to remove the male organs in fowls which are known
to be producing gametes of two kinds only. Further experiments

C. J. Bond 139

should also be undertaken showing the effect of partial removal of
the ovary in heterozygous hens, in order that the influence of re-
generation may be tested in the case of female as well as male
gamete-forming organs. There is, I think, no doubt from experimental
evidence that the ovary in the Fowl does undergo regeneration after
partial destruction or removal.

Thus the ovary was destroyed by the actual cautery in a hen
aged 1 year as far as could be seen with the naked eye wholly. On
examination 1 year and 9 months after the operation the ovary was
found to be fully regenerated and functionally active, a number of eggs
having been laid, see PI. V, fig. 10.

If these results are confirmed, that is to say, if it can be shown that
the effect of the stimulus to renewed growth which follows castration,
is to alter the character of or the proportionate rate of cell division in,
the cells which produce the male gametes then it will be necessary
to study these changes in cell division in detail, because it is possible
that certain results in breeding which seem to be inconsistent with
Mendelian expectation may owe their apparent anomaly to some change
in the rate of reproduction of gametes of different kinds in that particular
sex gland.

For instance, the normal relative proportion of male to female
chickens hatched at different times of the year varies, and this fact
suggests a seasonal difference in the relative production of male and
female zygote-forming sperm-cells.

[The explanation of Plates IV and V is to be found on the plates
themselves.]

MENDEL'S PRINCIPLES
OF HEREDITY

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Horticultural Institution. Tliird impression with additions. With
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By W. BATESON, M.A., F.R.S., V.M.H.

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which for surprising novelty and adventure is hardly to be excelled.' " — Morning Post

HEREDITY AND MEMORY

The Henry Sidgwick Memorial Lecture

delivered at Newnham College, Cambridge
9 November 1912

By JAMES WARD, Sc.D., Professor of Mental Philosophy in the University
of Cambridge, Author of The Realm of Ends or Pluralism and Theism.
Crown Svo. Paper covers, Is. net. Cloth, Is. Qd. net.

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Edward N. Wentworth. The Segregation of Fecundity Factors

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Heredity and Eugenics. By John M. Coulter, William E.
Castle, Edward M. East, William L. Tower, and Charles B.
Davenport. 312 pages, 8vo, cloth; 10s. net.

Five leading investigators, representing the University of Chicago, Harvard
University, and the Carnegie Institution of Washington, have contributed to this
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general reader the present position of evohition, experimental residts in heredity in
connection with both plants and animals, the enormous value of the practical
application of these laws in breeding, and human eugenics. Technicalities of
language have been avoided, and the result is an instructive and illuminating
presentation of the subject for readers untrained in biology as well as for students.

Contents : I. Recent Developments in Heredity and Evolution : General Introduction.

II. The Physical Basis of Heredity aud Evolution from the Cytological Standpoint (John
Merle Coulter, Professor and Head of the Department of Botany, the University of Chicago).

III. The Method of Evolution. IV. Heredity and Sex (William Ernest Castle, Professor
of Zoology, Harvard University). V. Inheritance in Higher Plants. VI. The Application
of Biological Principles to Plant Breeding (Edward Murray East, Assistant Professor of
Experimental Plant Morphology, Harvard University). VII. Recent Advances and the
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Organisms by Experimental Processes (William Lawrence Tower, Associate Professor of
Zoology, the University of Chicago). VIII. The Inheritance of Physical and Mental
Traits of Man aud their Application to Eugenics. IX. The Geography of Man in
Relation to Eugenics (Charles Benedict Davenport, Station for Experimental Evolution,
Carnegie Institution of Washington).

British 2Iedical Journal. Those who are desirous of arriving at an estimate of the
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"Heredity and Eugenics."

The Nation, New York. "Heredity and Eugenics" may be heartily recommended to

readers seeking, as beginners, to get in touch with the discussion of these subjects In

most of the lectures there is an admirable reserve, not to say skepticism, in the treatment
of large questions which the public is often misled to regard as already and finally settled.

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Kockefeller Institute for Medical Research. 23S pages, 12mo,

cloth ; 6s. net.

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that life may be unequivocally explained in physico-chemical terms. An affirmative
answer, he thinks, will necessitate a reconstruction of our social and ethical life on
a scientific basis. This volume is a popular presentation of the results of tlie
author's investigations, including his successful experiments in chemical fertilization.
The wide range of his discussion is seen in the following li.st of contents: I. The
Mechanistic Conception of Life. II. The Significance of Tropisms for Psychology.
III. Some Fundamental Facts and Conceptions concerning the Comparative
Physiology of the Central Nervous System. IV. Pattern Adaptation of Fishes
and the Mechanism of Vision. V. On Some Facts and Principles of Physiological
Morphology. VI. On the Nature of the Process of Fertilization. VII. On the
Nature of Formative Stimulation (Artificial Parthenogenesis). VIII. The Prevention
of the Death of the Egg through the Act of Fertilization. IX. The Role of Salts
in the Preservation of Life. X. Experimental Study of the Influence of Environment
on Animals.

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Volume III

FEBRUARY, 1914

No. 3

STUDIES OF INHERITANCE AND EVOLUTION
IN ORTHOPTERA I.

By ROBERT K. NABOURS.

LIBRARY
NEW YOKK
BOTANIC/i»l,

6AK»>fe.P»,

Paper 3. From the Zoological Laboratory, Kansas State
Agricultural College.

CONTENTS.

FAQE

I. Introduction 142

II. Material and Method 142

III. Analyses of gametic constitutions and the inheritance of colour patterns . 144

(1) An analysis and the inheritance of the colour patterns of forms of

the appearance of texanus, leuconotus, leucothorax and puncto-

femorata 144

(2) An analysis and the inheritance of the colour patterns of forms

of the appearance of luteolineatus and rufrolineatus . . 150

(3) The interbreeding of heterozygotes 155

(4) Further hybridization and the inbreeding of the heterozygotes . 157

(5) Results from the mating of heterozygotes with one or the other

of the parent forms 159

IV. Long and Short wingedness 160

(1) Long and short wingedness in nature 160

(2) Long and short wingedness in the breeding experiments . . 162
V. An examination of the location, arrangement and relations of the pigmental

elements in the colour patterns of the parent forms and their

hybrids 164

VI. Discussion 166

(1) The inheritance of the colour patterns 166

(2) The appearance of long and short wingedness .... 167

(3) Equivalence in the hybrids 168

(4) The "genotype conception " applied to the behaviour of these forms 168

VII. Conclusions 169

Bibliography ............ 170

Journ. of Gen. in 10

142 Inheritance and Evolution in Ortlioptera I

I. IXTKODUCTIUN.

The well-known cxpeiiments of Mendel, and the work of the Neo-
Mendelians with numerous plants and animals, show clearly that
surprisingly exact predictions of the results of breeding can be made,
provided the gametic constitutions of the parents are known, further-
more the gametic constitution of the parents can also be determined
by breeding analyses. That there is segregation, or alternativeness, in
gametogenesis which accounts for the familiar Mendelian ratios is
a generalization which now seems to be well established. However,
even in this matter there are some apparently important exceptions
which engender doubt in the minds of some persons.

The existence of unit characters, in the De Vriesian sense, does not
appear to have been as clearly demonstrated as that of alternative
inheritance, and if one may judge from expressions of opinion con-
cerning this matter, the interpretations are at great variance. Thus
one group of authors recognize characters in organisms that can be
replaced by other characters, when the proper crosses are made —
a clear recognition of separable and replaceable characters, which
are not necessarily unit characters (1, 3, 8) — while on the other side
there are those who believe that the organism as a whole is the only
unit and that there are no actual unit characters (9, 7).

In this paper is presented a preliminary account of an experimental
inquiry into the problems of inheritance and evolution, which is now
being carried on with several species of the grouse-locusts (Tetriginae)
of the genus Paratettix, Bolivar.

II. Material and method.

The Tetriginae are widely distributed, and are principally dis-
tinguished from other nearly related Orthoptera by the pronotum which
extends backwards over the body and wings, a character which varies
greatly among the different genera. The North American genera are
mostly geophilous, live on damp earth covered with algae, especially in
moist meadows and woods, and on the margins of ponds and streams (4).
The genus Paratettix, Bol., is distributed over a large part of the United
States and Mexico, and the species therein are mainly distinguished by
their striking colour patterns.

li. K. Nahouks 14:3

I have collected in the Miississippi Valley and on the Gulf Coastal
Plain of Louisiana and Texas nine distinct true breeding forms of this
genus which are being used in the breeding experiments (Plate VI,
tigs. 1 — 9). There seems to be uncertainty as to the taxonomic
position to be given these forms. It has been suggested that they
be called varieties and again that they be designated as biotypes, and,
with best reason, it seems to nie, that they be called species outright.
Pending further consideration of this matter, it seems sufficient for the
purposes of this paper to give them names descriptive of the colour
patterns, leaving to the future the question as to whether or not these
designations are to become the specific names of the forms. However,
the names will be tentatively submitted as though they are the specific
ones of the genus Paratettia-, Bol., the taxonomic position which I
believe will eventually be accorded them. The nine forms, all of them
new, except one, are as follows : Paratetti.r texanus, Hanc. (Plate VI,
fig. 1); F. leuconvtus, n. sp. (tig. 2); P. leucotltorux, n. sp. (fig. 3);
P. punctofemorata, n. sp. (fig. 4); P. lateolineatus, n. sp. (fig. 5);
P. rufrolineatus, n. sp. (fig. 6); P. melanothorax, n. sp. (fig. 7);
P. luteonotatus, n. sp. (fig. 8); and P. nifjronotutiis, n. sp. (tig. 9).
A more detailed taxonomic description of these and several other forms
used and something of their habits will be presented in a separate
account.

The characters used in this investigation have been the colour
patterns of the pronota and of the femora of the jumping legs (Plate
VI), and the length of the pronotum and wings, whether short, inter-
mediate, or long (Fig. 2, p. 161). The tinal colour pattern is clearly
indicated soon after the moult which ushers in the second instar, and
there are no perceptible changes in it during the remainder of the life
of the individual. The wing and pronotum lengths cannot be deter-
mined until the individual becomes adult, and are not in any way
correlated with the colour patterns. The forms and their hybrids
interbreed freely, and reciprocal crosses have invariably produced
identical results. All the females used in the experiments recorded
here were virgin, excepting the two heterozygous texanus-leucothoi-ax
females from nature used in the beginning of Exp. I (p. 145).

Glass cylinders, 8" x 12" and 9" x 15" respectively, set in pots of
earth and covered with 12- or 24-mesh pearl wire, seem to make the
most convenient cages (Fig. 1). It is found best to overlay the surface
of the earth in the cages with a very rich vegetable mould, or peat
substance. The smaller cages covered with the 12-inch mesh wire are

10—2

144 Inheritance and ^volution in Ortlwptera I

used as breeding cages. Soon after the young hatch, they are trans-
ferred by means of a damp camel hair brush to the larger cages covered
with the 24-mesh wire. The best food found consists of the green
scrapings (algae, lichens, etc.) from the long used pots holding hot-
house plants, though the various filamentous algae serve very well.
The mortality on the whole is great.

Fig. 1.

III. Analyses of gametic constitutions and the

INHERITANCE OF COLOUR PATTERNS.

(1) An analysis and the inheritance of the colour patterns of forms
of the appearance of texaiius, leuconotus, leucothoi'ax, and puncto-
femorata.

For convenience, the homozygous form.s an<l their gametes will be
designated by symbols (letters): texanus by .1, louconotus by B, leuco-
thorax by C, punctofemorata by D, luteolineatus by E, rufrolineatus
by F, melanothorax by G, luteonotatus by H, and nigronotatus by /,

R. K. Nabours 145

Any two of these letters placed side by side will rejjresent the hetero-
zygous, or hybrid zygote, or individual, produced by the union of the
homozygous gametes represented by the respective letters.

Early in September, 1908, specimens of the general appearance of
texanus, leuconotus, leucothorax, and punctofemorata (Plate VI, figs.
1 — 4), were obtained in the vicinity of Houston, Texas. At that time
nothing whatever was known of their composition, and the females were
not virgin. All the forms were placed in a tin bucket and taken to
Chicago. One specimen, a male of the appearance of leuconotus
(Plate VI, fig. 2), two specimens, females, of the appearance of leuco-
thorax (Plate VI, fig. 3), and several males and females of texanus
and punctofemorata survived (Plate VI, figs. 1 and 4).

Experiment I. On September 12th, the male of the appearance
of leuconotus was mated to the two females of the appearance of
leucothorax (Table I, parents). The progeny which hatched in
November and December grew slowly and after great mortality
matured in the following March, and showed in F^ five types as
follows : 5 of the appearance of leuconotus, 6 leucothorax, 6 inter-
mediate between leuconotus and leucothorax, 12 texanus, and 3 puncto-
femorata (Table I, Fi). The inbreeding of these Fi types, each type
to itself, in the F.^ generation was as follows : two pairs of the leuconotus
tjrpe were mated in separate cages and gave Fo progeny as follows : in
cage (a) 34 leuconotus (B) and texanus-leuconotus {AB) : 14 texanus
(A), and in cage (6) 24 leuconotus {B) and texanus-leuconotus (AB) :

7 texanus (A); the total numbers for the two cages being 59 leuconotus
(B) and texanus-leuconotus (AB) : 21 texanus (A), with the expectation
of 60 : 20, respectively. The F^ leuconotus-leucothorax (BC) hetero-
zygotes were inbred, as a groujj culture, and gave in F,,, 66 leuconotus;
136 leuconotus-leucothorax (BC): 58 leucothorax, the expectation being
65 : 130 : 65, respectively. The Fi leucothorax type inbred, three pairs
in separate cages, gave progeny in Fo as follows : in cage (a) 43 leuco-
thorax and texanus-leucothorax (AC) : 7 texanus (A); in cage (b)

8 leucothorax (C) and texanus-leucothorax (AC) : 10 texanus (A);
and in cage (c) 66 leucothorax (G) and texanus-leucothorax (AC) :
15 texanus (A); the total numbers from the three matings being
117 leucothorax (C) and texanus-leucothorax (AC) : 32 texanus (A),
with the expectation of 11175 : 3725, respectively. One of the F^
punctofemorata type, a male, was mated to a texanus female, and they
gave in F„, 9 of the appearance of punctofemorata : 5 texanus. This
result shows at least the texanus-punctofemorata (AD) heterozygous

140 Jnlieritance and Erolufioii in Ortlioptcrd T

co!istitutioii of the F^ individuals of the appearance of punctofemorata.
The F^ texanus bred true for five generations when the culture was
terminated. For a graphic illustration of all these F., results compare
Table I, F.„ and diagram 1, jf, (p. 151).

At this time, Spring 1909, I was not able to distinguish the pure
leuconotus from the hybrid leuconotus-texanus, but since then I have
learned to distinguish between them. On careful examination, the

TABLE

I.

E:

spectation

3

G

3

Actual Numliers

')

4

3

21

•

•

•

•

B

BC

C

C

1

1

1

Lxpeclation

28-25

56-5

28-25

Actual Numbers

8

29

53

31

11

•

•

•

•

•

B

B

1

BC

C

J

C

Expectation

7'o

15

7-5

Actual Numbers

3

21

1

10

17

3

y

2

r 1

v

1

•

t t

•

•

•

•

•

t

A

AB B

B

BC

C

C

AC

A

1

1

1

1

1

'i

1

i

1

Expectation

231-25

462-5

231-2;

Actual Numbers

107

977

251

452

1

222

r

454
1

29

•

• t

•

•

•

•

•

•

A

AB B

B

BC

C

C

AC

A

L

—

1 -"

1

1

-J

L.

1

1
1

I
r— '

E.\pectation

20

60

C5

130

65

1 1 1 -75

37-25

Actu.il Numbers

21

59

G6

136

58

117
1 ^

32

r"

n

•

• •

•

t

•

•

•

t

A

1

AB B

1

B

1

BC

C

1

C

1

AC

A

1

1

1

1

1

1

-r
j

1

5

G

6

12

3

t

•

•

•

•

AB

BC

AC

A

AD

1

•

1

1

X

•

1

Pa

AB

AC

1 6

from

2

9 s from

the

field

the field

not

\-irgiu

A^

= texainis.

C =

= leacothora.i:

B

= leuconotus.

D =

- pa DctofemoraUt .

F,

F,

R. K. Nabours 147

general outlines of the pattern of texanus can be clearly seen in the
heterozygote. I am not able to distinguish with certainty between any
of the other pure forms and the hybrids between them and texanus ;
that is, as ordinarily considered, texanus is recessive to all the other
species.

An analysis of the Fo results just described, with their close approxi-
mations to the expected ratios of alternative inheritance, shows that
the Fi leuconotus were heterozygous, having the gametic constitution
AB (allowing A and B to represent the gametes of texanus and leuco-
notus, respectively). The leucothorax in the same way were also,
heterozygous, having the composition AC {C representing leucothorax),
and the behaviour of the midtype strikingly points to its heterozygous
leuconotus-leucothorax {BV) character. From this behaviour it seems
that we may conclude that the parent male from nature was a texanus-
leucouotus heterozygote of the gametic constitution AB, and the
females (at least one of them) were texanus-leucothorax {AC) hetero-
zygotes and that some of the eggs of at least one of these females had
been fertilized by a pure punctofemorata (Z)), or heterozygous texanus-
punctofemorata {AD) male, and his D gametes meeting her A gametes
made the heterozygous texanus-punctofemorata {AD) specimens of the
superficial appearance of punctofemorata (Table I, Fi, and diagram 1 F-,).

F-, results from inbreeding of the F„ forms of this series. The
leuconotus-leucothorax {BC) inbred, as a group culture, gave in F^
251 leuconotus : 452 leuconotus-leucothorax (BC) : 222 leucothorax,
the expectation being 281-25 : 4625 : 231-25, respectively. The F,
generation leuconotus which came from the inbreeding of F^ leuconotus-
leucothorax bred true for three generations and then the culture was
destroyed. The F., leucothorax which came from the inbreeding of
this F-i leuconotus-leucothorax midtype were carried two generations
and bred true. The F, forms of the general appearance of leuconotus
were inbred in four cages as follows : (a) 1 male x 1 female, (6) 1
male x 1 female, (c) 2 males x 2 females, {d) 2 males x 2 females. Pair
(a) gave 10 leuconotus : 3 texanus; pair {h) gave 275 all leuconotus;
pair (c) gave 194 leuconotus : 47 texanus; and pair {d) gave 498 leuco-
notus : 57 texanus. These results are represented graphically in Table I,
F^, as a group culture, as unfortunately they essentially were, because
the main attention at this time was being given to the behaviour of
the leuconotus-leucothorax midtype and its products, leuconotus and
leucothorax, and to the finding of the proper food and other living
conditions. However, the results are suggestive; for in cage («)

148 Inlieritanee find Evolution in Orthoptera I

tli<mgh the nmnbers are small, the 10 : 3 ratio suggests that the
parents were texamis-leiiconotus (AB) heterozygotes, since this is a
close approximation to the expectation of 975 £ and AB -.^^^ A. In
cage (6) one of the parents could have been of the constitution AB and
the other pure B, or botii could have been pure B, because at that time,
as has already been noted, in making records no distinction was made
between the heterozygous texanus-leuconotus {AB) and the homo-
zygous leuconotus {B). The specimens have been kept preserved in
alcohol and although the general outlines of the patterns are still
distinct they are not clear enough to permit the fine distinction which
would be necessary for selecting the homozygous leuconotus {B) from
the hetei-ozygous texanus-leuconotus {AB), as can be done now with
the live specimens.

It is obvious that nothing further than that they are AB and B can
be ascertained concerning the composition of the parents in cages (c)
and {d), though the ratio of 194:47 in cage (c) is suggestive of the
expectation of 18075 : 60'25, if all four of the individuals had been
texanus-leuconotus {AB) heterozygotes. Although the matings were
made in such a way as to make the numerical ratios of no value, yet
it is to be noted that no unexpected individuals appeared — all were
texanus {A), texanus-leuconotus {AB), or leuconotus {B).

The F.:, impure forms of the appearance of leucothorax were inbred
in the same way as were the F,, '\m\mre leuconotus, i.e., essentially as
a group culture. As in the case of the homozygous leuconotus and the
heterozygous texanus-leuconotus, I was not able to distinguish between
the homozygous leucothorax and the heterozygous texanus-leucothorax.
Four group matings were made as follows : (a) 1 male x 1 female ;
(6) 2 males x 2 females ; (c) 2 males x 2 females, and {d) 2 males x
2 females. The F-,, results from these matings were as follows : from
(a) 259 leucothorax; {b) 65 leucothorax : 11 texanus; (c) 103 leuco-
thorax : 15 texanus; and {d) 27 leucothorax : 3 texanus. If none of the
leucothorax F^ individuals from (a) had been bred further, it would not
be known whether the F., parents were both homozygous leucothorax {C),
or one of them homozygous (C) and the other heterozygous texanus-
leucothorax {AG), but 2 males X 2 females were bred and gave in Fi,
after great mortality due to drought in the cage, 9 leucothorax : 2
texanus; thus showing one of the parents in cage (a), F.,, to have
been undoubtedly heterozygous texanus-leucothorax {AC). The in-
dividuals from the other cages were not bred further. It is here
again obvious that the composition nf each of the paivnts in these

R K. Nabours 149

cages cannot be ascertained by the appearance of their F^ offspring,
because 2 males x 2 females were used in each instance. However,
no unexpected types appeared, and we do know that the parents
were of the two compositions, C and AC, and no other than these.
The results from the inbreeding of these F., impure leucothorax are
shown as a group in Table I, F,, and in more detail in diagram
1, F,.

A group culture of 2 males x 2 females was made from the Fo
texanus which had come from the ^i impure leuconotus and they
gave only texanus in F^ when the culture was terminated. Similarly,
a group culture was made of the F„ texanus which had come from the
Fi impure leucothorax and they bred true in F^ when they were also
destroyed.

Fi results from the separate inbreeding of the F^ forms of this
series. Several of the F-^ leuconotus-leucothorax {BC) were inbi-ed,
but on account of poor attention, the mortality of the F^ progeny
was very great. The record of those reaching maturity was 10
leuconotus : 17 leuconotus-leucothorax : 3 leucothorax (Table I, F^).

From {d), of the texanus-leuconotus {AB) F^ group culture, two
pairs, a pair each in separate cages, of the appearance of leuconotus were
bred. After great mortality the F^ result in (o) was 8 leuconotus : 3
texanus, and the pair in (h), gave 13 all leuconotus. These results
added together are shown in Table I, F^, as a group culture, giving
a 21 : 3 ratio. It appears that the parents in cage (a) were texanus-
leuconotus (AB) heterozygotes, and that the cage (6) parents were both
either homozygous leuconotus {B), or that one of them was homozygous
and the other heterozygous texanus-leuconotus {AB). However, when
a pair from cage (6) were bred, they gave in F^, 8, all leuconotus in
appearance, and this closed the impure leuconotus part of the experi-
ment (Table I, F^ and F,).

From the cage (a) i^s progeny, in the impure leucothorax series of
this experiment, two males and two females of the appearance of leuco-
thorax were mated as a group culture, and they gave in Fi, 9 leucothorax :
3 texanus. This result, though small in numbers, shows quite clearly
that the pair of the appearance of leucothorax taken from the (o) F^
culture and bred to make (a) F^ 259 all leucothorax in ajipearance,
were not both of them homozygous leucothorax {G), nor were they both
heterozygous texanus-leucothorax {AC), but that one of them was a
homozygous leucothorax (0), and the other a heterozygous texanus-
leucothorax {AC), a matter to which attention is given above. The

150 Tnheritancf and Evolution in Orthoptera I

leucothorax progeny from this F^ culture bred true, when inbred, lor
two generations, giving 11 leucothorax in F, and 21 leucothorax in F,i
(Table l,F,,F,sind F,).

A pair of texanus from the Fi impure leuconotus culture,
when bred, gave 100 °/^ texanus, as did a similar pair from the
same generation of the impure leucothorax culture. Also a texanus
male from the F^ texanus-leuconotus culture was mated to a texanus
female from the F^ texanus-leucothorax culture and they gave 100 7o
texanus.

The Fi leuconotus-leucothorax (56') were inbred and gave, in
F^, 29 leuconotus : 53 leuconotus-leucothorax \BG) : 31 leucothorax,
the expectation being 28-25 : 56-5 : 28-25, respectively. (Table I, F^.)

The Fr^ leuconotus-leucothorax {BC) were again inbred and gave
in F,i, 5 leuconotus : 4 leuconotus-leucothorax (BC) : 3 leuconotus
(Table I, F^). With this result the experiment was closed.

Allowing the letters A, B,C, and D, as already suggested, to indicate
the homozygous texanus, leuconotus, leucothorax, and punctofemorata,
respectively, diagram 1, with Table I, will show at a glance the inherit-
ance behaviour in this experiment.

(2) An analysis and the inheritance of the colour patterns of forms
of the appearance o/'luteolineatus and rufrolineatus.

Experiment II. Early in September, 1909, among several specimens
secured in the vicinity of Many, Louisiana, were a male of the appear-
ance of rufrolineatus (F) (Plate VI, fig. 6) and an innnature female of
the appearance of luteolineatus(£') (Plate VI, fig. 5). When the female
became adult this male of the appearance of rufrolineatus was mated
to her, and their progeny wiiich hatched in November, after great
mortality, matured the following March and showed four types in Fj
as follows : 4 of the appearance of rufrolineatus : 2 intermediate between
rufrolineatus and luteolineatus : 2 of the appearance of Inteolineatus :
2 texanus (Table II, (a), Fi). These F^ progeny were bred further
as follows : (a) a pair of the appearance of rufrolineatus were inbred
and gave in F^ 43 of the apf)earance of the parents (rufrolineatus) : 13
texanus. This shows clearly that the Fi individuals of the appear-
ance of rufrolineatus were really texanus-rufrolineatiis (AF) hetero-
zygotes. This fact, the visibly clear luteolineatus-rufrolineatus (EF)
heterozygous cliaracter of two others of the Fi progeny and the fact
that still two others were the always homozygous texanus (A), without
considering the two inilividuals of the appearance of luteolineatus wliieh

R K. Naboitrs

151

s <

<

o

a

E

«o CD

^ dnoia " " dnoja "" dnojii ^ ™ dnoja
2 '>° 5 OQ

S 00

00 CO

O
<

<

S

o

CD <(

152 Inheritance and Evolution in Orthoptera T

^

M nq • O J

CM <N

- ~ ^ • ^ -1
^ •>] • O

CC (35 (
CM f>l

■o -

°0 -t — ,^
CM Oq • Cj

_0 1- » "-

o5

HO

iij ^

"TDUJ

SS^"^

»c

W

i-:i

o

m

I-

<;

S ;2 •03

-co ^ A llJ
« fO CD ■

g a I

= c
& S

^ (ij qj ~

< OQ O liJ IJ-

Z^»<^

i..

I • <-l

■•^

o+.£-=

<u .

♦o

■Z%^

.2-3^

II

u

a» ,_

o ^

^ ce

<

<

R. K. Nabours 153

were not bred, all show that the parents fi-oni nature were really texa-
nus-rufrolineatiis {AF) and texanus-luteolineatus (AE) heterozygotes,
respectively. Therefore, the F^ progeny should read, texanus-rufro-
lineatus (AF) 4 : luteolineatus-rufrolineatus (EF) 2 : texanus-luteo-
lineatus (AE)2 : texanus 2, the expectation being 2'25 of each (Table
11 (a), F,).

(b) One of the luteolineatus-rufrolineatus (EF) Fi males (Table
II (o) Fi) was mated to an extracted homozygous leuconotus (B) iemale,
from a culture whose parents were both leuconotus-leucothorax (BC)
heterozygote (Table I, F^, B) and they gave F^ progeny as follows:
leuconotus-rufrolineatus (BF) 31 : leuconotus-luteolineatus (BE) 36,
the expectation being 33-5 of each kind (Table II (b), Fj). This
result proves unmistakably the luteolineatus-rufrolineatus (EF) hete-
rozygous character of the male parent, supplementing the evidence
which the intermediate colour of his stripe had given.

The inbreeding of these leuconotus-rufi'olineatus (BF) and leucd-
notus-luteolineatus (BE) F^ individuals, respectively, gave F.. results
as follows : two pairs of the leuconotus-rufrolineatus (BF) individuals
were mated, one pair in each of two cages. The pair in cage (a) gave
the following F. progeny: leuconotus 27 : leuconotus-rufi-olineatus (BF)
69 : rufrolineatus 29; and the pair in cage (b) gave leuconotus 24:
leuconotus-rufrolineatus (BF)71 : rufrolineatus 30; the two pairs giving
totals of 51 leuconotus (B) : 140 leuconotus rufrolineatus (BF) : 59 rufro-
lineatus (F), with the expectation of 62'25 : 1245 : 62'25, respectively
(Table II (b), F,).

One of these F2 rufrolineatus males was mated to a sister leuc(jnotus-
rufrolineatus female and they gave in F, 6 leuconotus-rufrolineatus :
5 rufrolineatus, the expectation being 5'5 of each. A pair of the F„
extracted rufrolineatus were inbred and gave in F, 100 °/^ rufrolineatus.
These were again inbred, as a group culture, and gave in F^ 100 °/^
rufrolineatus.

Two pairs of the F^ leucondtus-luteolineatus (BE) were mated, one
pair in each of two cages. Pair (a) gave 32 leuconotus : 53 leuconotus-
luteolineatus (BE) : 17 luteolineatus, and pair- (b) produced 33 leuco-
notus : 48 leuconotus-luteolineatus : 20 luteolineatus, both cages giving
totals of 65 leuconotus : 101 leuconotus-luteolineatus : 37 luteolineatus,
the expectation being 50'75 : lOl'o : 5075, respectively (Table II (b),

Three males and three females of these F.^ leuconotus-luteolineatus
(BE) individuals were inbred, as a group culture, under very unfavourable

154 Inheritance and Erohition In Orlhoptera 1

conditions and they gave in F^ !S louconotus : 19 leiiconoUis-luteo-
lineatus : 6 luteolineatus, the expectation being 8'25 : 165 : 8'25,
respectively. One of the luteolineatus F^ males (Table II (t), F.^
was mated to a virgin female of the same apparent constitution but
whose pedigree is not known, and they gave in F-^ 100 7o luteolineatus.

(c) The other luteolineatus-rufrolineatus {EF) heterozygous male
(Table II (a), F^ was mated to an extracted homozygous leucothorax
(C) female from a culture whose parents were both leuconotus-leuco-
thorax (5C) heterozygous individuals (Table I, i^^, C). (This female
was a sister of the leucontitus female parent in mating (b) »{ this
experiment.) They produced the following F^ progeny : leucothorax-
rufrolineatus {CF) 17 : leucothorax-luteolineatus 21, the expectation
being 19 of each kind (Table II (c), F,\ The behaviour of this EF
male when mated to a homozygous leucothorax female was exactly the
same as that of his brother when mated to a homozygous leuconotus
female. Both behaviours reveal the constitution of the males, as well
also as that of the females.

The inbreeding of these F^ leucothorax-rufrolineatus (GF) and
leucothorax-luteolineatus (GE) individuals, respectively, gave F^ results
as follows : two pairs of leucothorax-rufrolineatus (GF) were inbred, a
pair in each of two cages. The pair in cage (a) produced 7 leuco-
thorax : 13 leucothorax-rufrolineatus (GF) : 4 rufrolineatus, and the
jjair in cage (b) gave 17 leucothorax : 48 leucothorax-rufi'olineatus
{GF) : 25 rufrolineatus ; both pairs giving a total of 24 leucothorax : 61
leucothorax-rufrolineatus (GF) : 29 rufrolineatus, the expectation being
28-5 : 57 : 28-5, respectively (Table II (c), F.^

The leucothorax-luteolineatus {GE) were inbred, one jjaii' in each
of two cages. Pair (a) produced 12 leucothorax : 12 leucothorax-luteo-
lineatus {GE):S luteolineatus, and pair (b) gave 10 leucothorax : 41
leucothorax-luteolineatus (GE): 16 luteolineatus; both pairs producing
totals of 22 leucothorax : 53 leucothorax-luteolineatus (GE) : 19 luteo-
lineatus, with the expectation of 23'5 : 47 : 235, respectively (Table
II (c), F.;).

These F.. leucothorax-luteolineatus (GE) were inbred under excep-
tionall}' unfavourable circumstances, 1 male x 1 female, and they gave
in F;, 8 leucothorax : 6 leucothorax-luteolineatus : 3 luteolineatus, the
expectation being 4'25 : 8"5 : 425, respectively. A pair of these F^^
leucothorax-luteolineatus individuals were inbred, and after great
mortality, gave in F^, 4 leucothoiax : 1 leucothorax-luteolineatus (GE):
2 luteolineatus, the expectation being 1'75 : 3'o : r7o, respectively.

R. K. Nabours

155

(3) The interbreeding of heterozijgotes.

Experiment III. All the crosses of heterozygous individuals with
homozj'gous ones and the inbreeding of heterozygous ones had given
results that could hardly be interpreted in any other way than by
assuming that the heterozygous, or hybrid, individuals gave gametes
alternatively, one representing the male j)arent and the other the
female parent. Not a single exception to this rule had been noted.
A few crosses had been made between different heterozygotes from
nature whose constitutions were not ascertained till their Fi and F.^
progeny gave the clue (Exp. I, Table I, and Exp. II (a), (6), (c),
Table II (a), (6), (c)). At this time it seemed that some crosses
ought to be made between ditfereut hybrids of known constitutions ;
so the following crosses were made simultaneously :

((f) A leucothorax-luteolineatus {CE) male from Exp. II (c), F^
(Table II (c), -fj) and a leuconotus-leucothorax {BC) female from
Exp. Ill (c), Fi (Table IV (e), F^) were crossed, and they gave in the
F^ generation {F„ Table III («)):

Actual Numbers ...

Leucouotus-
Leucothorax leucothorax
(C) [BC

9 9

Leucothorax-
luteolineatus

7

Leucouotus-
luteolineatus
{BE)

13

Expectation

9 -.5 9o

9-5

9-5

TABLE III

Expectation
Actual Numbers
^1

9o
: 9

•
C

:

9-5
9
•
BC

1

9-.". 9-.5
7 13
• •

CE BE

1

4.5

•
CE

1

45
36

•
BC

1

45

44

•

BF

45

48

EF

1

Parents

CE

I
X

BC

1
CF

$ • Parci
BE

From Tab!

II (C) (i^ij

le

1

(a

Fi-om Table From

\X{C){F,) II (C

Table

From Table

II (b) {F,}

Expectation
Actual Numliers

21-5

18

f

BE

L

■2\-b

23

•

CE

21 -.5 21-5
i\ 24
t •
BC E

10
12

•
BC

1

10
11
•
BE

10
7
•
CI

10 0
10 1

• t

El BEI

1

Parents

CE

1

BE

CE

1
X

9 • Parei
Bl

From Table
11(C) Z',

From Tabic

II(b)(i^,)

From Table
II(C)(/',)

Group
culture

/',

(c)

(e)

B = leticonotus. C = leucothorax. E=luteolii>,e<-itug.

F = rii/ruluieiittis. I = iiigronoOxtus.

l.')() Inheritance and Evolution In Orthoptera I

{b) A leucothorax-rufrolineatus {GF) male from Exp. II (c), F^
(Table II (c), F^ was mated to a leuconotus-luteolineatus {BE) femak-
from Exp. II {b), F, (Table II (6), F,) and the re.sulting F^ progeny
were as follows {F„ Table III (b)) :

Lcucothorax-
luteolineatus

Leuoonotus-
leucothorax
(i'Cl

I.euconotus-

rufrolineatus

iBF)

Luteolineatus-
rurrolineatus
(EF)

Actual Numbers ...

52

36

41

48

Expectation

45

45

45

45

(c) A leucothorax-luteolineatus (GE) male from Exp. II (c), F^
(Table II (c), F^) was crossed with a leuconotus-luteolineatus {BE)
female fn)m Exp. II {b), F, (Table II (b), F,) and they gave in the
Fi generation (Table III (c), F,):

Actual Numbers
Expectation

Leuconotus-
luteolineatus
iBE)

Leucothorax-
luteolineatus
iCB)

Leuconotus-

leucothorax

(BO

Lut«olilieatus
(B)

18

23

21

24

21-5

21-5

21-5

21 -.5

{d) A leuconotus-luteolineatus {BE) male from Exp. II (b), F-. and
a female leuconotus-leucothorax (56*) from Exp. Ill (e), F^ were mated
;\\n] the}' gave in the F^ generation the following progeny :

Leuconotus
(B)

Leuconotus-
leucothorax
(BC)

Leucothorax.

luteolineatus

(VE)

Leuconotus-
luteolineatus
(££)

Actual Numbers ...

7

7

9

fi

Expectation

7-25

7-25

7-25

7-25

This experiment, (rf), was carried on subsei|ueutly to (c) which is the next experiment
in this series to be described.)

The results from these crosses gave nothing new. They can be
accounted for clearly by assuming that the parent heterozygotes gave
gametes alternatively for each parent type of which they were composed
and that in fertilization these gametes met by chance. Considering
the small numbers, it seems that the approximations to the expectations
of alternative inheritance are fairly close. However, the next cross (e)
in this series is not so regular.

(e) A leucothorax-luteolineatus {GE) male from Exp. II (c), F^
(Table II (c), F^) was mated to a leuconotus-nigronotatus {BI) female
from a cross between an e.xtracted leuconotus {B) male and a nigi'o-
notatus (Plate VI, Fig. 9) female from nature. This leuconotus-nigro-
notatus female had the exact appearance of many other heterozygous

Luteolineatus-
nigronotatus
(EI)

Leuconotns-
luteolineatus-
ni^onotatus
[BED

10

1

R. K. Nabours 157

leucoiK^tus-nigronotatus (BI) individuals thab have since been produced.
Their F, progeny was as follows (Table III (e), F^) :

Leuconotus- Leuconotus- Leucothorax-
leucothorax luteolineatus nigronotatus
{BO (BE) [CI)

Actual Numbers 12 11 7

Expectation 12-25 12-25 12-25 12-25 0 F,

This result is perfectly regular in every respect, except tor the
appearance of the leuconotus-luteolineatus-nigronotatus (BEI) in-
dividual. This specimen was observed during its second instar when
its characters were perfectly distinct, and it was kept under clo.se
observation until it became adult. The pattern was distinct at all
times, but was strikingly clear just after moults. Unfortunately it
escaped while the culture was being transferred from the University
of Chicago to the Kansas State Agricultural College, at Manhattan,
in September, 1910. It seems that the production of this aberrant
individual may be accounted for by assuming that the leuconotus-
nigronotatus {BI) female parent gave at least one gamete containing
the factors for the patterns of both of her parents and that this double
character gamete was fertilized by one of the luteolineatus (E) gametes
which came from the leucothorax-luteolineatus (GE) male'.

(4) Further hyhidization and inbreeding of the resulting hetero-
zygotes.

E.Tperimeiit IV. The following are additional data concerning the
simple crossing of forms where the composition of one of the parents
was known and that of the other parent unknown.

(a) A male nigronotatus (Plate VI, fig. 9), from nature, whose
pedigree was not known, and a female leuconotus, fiom the F.^ genera-
tion of a pure leuconotus culture, were crossed, and they gave in Fi ten
adults, all leuconotus-nigronotatus(i^/) heterozygotes (Table IV (a), F^).
A pair of these were inbred and gave in F., three types as follows
(Table IV (a), F,) -.

Leuconotus

Leuconotus nigronotatus Nigronotatus
IB) {BI) (/)

Actual Numbers ... 3 13 5

Expectation ... 5-25 10-5 5-25

' In subsequent experiments — the production of an indindual with the patterns of
three of its ancestors combined — has been repeated four times with other combinations of
patterns. I now have leuconotus-rufrolineatus-melauothorax (BFG), leucothorax-rufro-
lineatus-melanothorax (CFG), leueonotus-leucothorax-melanothorax [BCG), and rufro-
lineatus-melanothorax-leuconotatus (FGJ) (the leuconotatus (J), a new form to be
described later). These have now been bred for some time, some of them to the F^
generation. TLeir inheritance behaviour will be reported soon in another account.
Journ. of Gen. iii 11

158 Inheritance (iiid Evolution in Ort/ioptini /

These were bred further as follows: a male leuconotns was mated to
a sister leuconotus-nigronotatus (Bl) female, and they gave in F^ the
following progeny :

l—" i

Leuconotus

Leuconotus-
nigronotatus
(Bl)

Actual N

umbers ... 4

6

Fz

Expectat

ion ... 5

5

TABLE IV.

Expectation
Actual Number;
F-.

0-25
s 5

•
1

10-5 5-25 4-2ri S-.-)
13 3 4 .s

• • e •

Bl B B BH

1 1

4-2.-5
5

•
H

1

5

•
B

1

17

!.■)

•

BC

25

•

BC

1

7-5
8

• F:

C

i

F,

1 1
10 15
t •

Bl BH

1 1

F,

Parents

r"

1

1 1
B B

1

?•

H

1
B

9 f Parents
C

Fr

om Group
Culture

Extracted Extracted
B Table I B Table I

Group
Culture

Extracted
B Table I

Extracted
C Table I

(a) (b)

(c;

1

B =

= lenconoiug. H =

luteonoUUns.

C =

= leucothorax. 1 = /

'ligroaotatiis.

These i^^ leuconotus-nigronotatus {Bl) were inbred, 1 male x 2 females,
and they gave in i^j the three following types :

conotus
(£)
9

Leuconotu.s-
nigronotatu?
(Bl)

30

Nigronotatus
(/)
13

13

26

13

Actual Numbers ... 9 30 13 t\

Expectation

{h) A male leuconotus {B), from the F„ generatinn of a pure
leuconotus culture, and a female luteonotatus {H), whose parents had
been taken from nature, were mated, and their F^ progeny were all
distinctly marked leuconotus-luteonotatus heterozygotes (Table IV (6),
F-^. Two males and two females of these were inbred and, after great
mortality, they gave in Fo the following progeny (Table IV (6), F.,):

Leuconotus-
Leuconotus luteonotatus Luteonotatus
(if) (BH) (H)

Actual Numbers ... 4 8 5

Expectation ... 425 S-.'j 4-2.^ F..

The.se results, as in the other cases (E.\p. II (a), (b), (c)), give the
clue to the composition of the unknown first parent in each case,
proving them to have been homozygous nigronotatus (7) and luteo-
notatus (H), respectively.

R. K. Nabours

159

(c) In order to try out more completely the behaviour of the
heterozygotes derived from the crossing oi' leuconotus and leucothorax,
a leuconotus male from the second generation of an extracted leuconotus
culture (Exp. I, F3) was mated to a leucothorax female from the second
generation of an extracted leucothorax culture (Exp. I, Fs). The F^
result was twenty-five adults, all leuconotus-leucothorax (BG). These
leuconotus-leucothorax heterozygotes were inbred and gave in F. the
following progeny (Table IV (c), Fj) :

Actual Numbers
Expectation

Leuconotus

(£)

5

7-0

Leuconotus-
leucothorax
{BV)

17
15

Leucothorax

i'-i

7-5

This result is in perfect accord with the other results, showing that
the first parents were homozygous, and that the Fi heterozygotes
behaved in a regular Mendelian manner just as did the exactly similar
leuconotus-leucothorax (BG) heterozygotes in Exp. I.

(5) Results from the matinf/ of heterozygous individuals -with one or
the other of their homozygous parent forms.

Experiment V. (a) A leuconotus-leucothorax (BG) male fi-om
Exp. I, Fs (Table I, F,) was mated to a leuconotus (B) female sister,
and they gave in F^ (F^^) the fijllowing results (Table V):

Actual Numbers
Expectation

Leuconotus-
leucothorax
(BC)

20

20

Leuconotus
(C)

20
20

Fi (Fi)

Expectation
Actual Numbers

Parents (Fs)

20

20

•

BC

TABLE V.

20
20

•
B

I

I

•

BC

BC Table I

(Fs)

(a)

B

Extract
TaLile I

(^3)

67 -.5
66

C

I

c

E.xlracL
Table I

B = leuconotus.
C = leucothorax.

(b)

67-.5
69

BC

I

•

BC
Table I

(^3)

^1(^4)

Parents (F3)

11—2

I()(> Inheritaiirc and involution in Ofthopt(i(( I

(b) A leucoiiotus-lcucothurax iniilc tVom Exp. I, Fj (Table I, F3)
and a sister Icucothoiax (C) were mated and their F, (Ft) progeny were
as follows :

LeiK^oiiotus-

leucotliorax Leucothorax

(BC) (C)

Actual Numbers ... 69 66 ^i (t't)

Expectation ... 67 -5 67-5

These results are typically Mendulian, and close the report to be
made at this titne on the inheritance of colour patterns.

IV. Long and shokt wingedness.

(1) Long and sliurt wingedness in Nature.

It is a matter of common observation that in many species of the
Acridiidae and Gryllidae there is dimorphism or polymorphism in the
length of the wings — some of the members bearing short and others
long wings. (In the Tetriginae, unless otherwise indicated, the word
wingedness refers to pronotum also.) The differences in some cases
have been considered by systematists sufficient to justify the giving
of different varietal names to the two forms (e.g. Tettigidea parvipennis,
Morse, and Tettigidea. parvipennis peninita, Morse, the short and long
winged forms of T. pennatus) (4).

The studies of the difference in lengths of the wings in the Acridiidae
have been, so far as I can ascertain, confined to the field observations
and to the examination of collections in museums. However, in the
Gryllidae, Lutz (1907) has made observations concerning this pheno-
menon of long and short wingedness (pronotum not considered) in a
breeding experiment with Gryllus sp. His results brought him to the
conclusion that the length of the wings of the species with which he
worked was not conditioned by heredity, but by the environmental
conditions under which the individuals grew to maturity (6).

In the Tetriginae, the differences in the length of the wings and
pronota are usually, though not invariably, dimorjjhie ; several varia-
tions from the long and short winged forms have been found, and some
individuals were strictly intermediate between the two extremes. It is
the usual occurrence to find a long pronotum with long wings and a
short pronotum with short wings, but a few \'ariations from this rule
have occurred as follows : in a few instances long wings have accom-
panied a short pronotum, and one individual exhibited a short pronotum
with one wing long and the other short (the various types are shown in
Fig. 2).

R. K. Nabours

161

Fig. 2.

1, 2, 3, Brotliers from the same cage.
4, 5, Brothers.

6, Short pronotum, both wings long.

7, Short pronotum, one wing long.

8, 9, Sisters.

All on the same scale.

162 InherUan.ce and Evolution in Orthoptera I

111 the vicinity of Chicago, the genus Tettigidea is found to be about
equally divided between the long and short winged individuals. They
are almost strictly dimorphic. The genus Tettix exhibits polymorphism
in respect to this character, running all the way from extreme short to
extreme long wingedness. The shorter winged individuals predominate
in this genus, while the extreme long winged ones are rare. In the
genus Furatettvr, Bol., under more particular consideration in this
jJaper, long wingedness is the rule in the Chicago region — only two
short winged individuals having been observed among many hundreds.
In Arkansas, a few short winged individuals were found among several
hundred long winged ones, while in Louisiana and Southern Texas, the
two forms, practically without intermediates, existed in about equal
proportions. In the North, in nature, only one generation a year of
any of the species is produced and the growth period is normally in
the late spring and early summer when the adult stage is reached
quickly, while in Louisiana and Texas Paratettix produces two or three
generations a year, and the growth period continues practically through-
out the year, with the optimum in the spring and early summer.

(2) Lone/ avd short tuivgedness in tlie breeding experiments.

The observation on the occurrence of lono' and short winored forms
during the breeding experiments will be reported at this time only for
the individuals maturing in the inheritance of the colour pattern in
Exp. I (Table I). In that experiment the wing length of the individuals
was not considered just as in the following account the colour patterns
are not considered, there being no apparent correlation between the
length of the wings and the pattern.

Referring to the diagram (diagram II), the first mating was made
in September 1908 between one short winged male and two short
winged females. At least one of the females was not virgin. The
Fx generation which hatched in December matured after great mortality
the following March and April in these proportions :

Long winged 17 : Short Winged 15 (i^i).

The short winged individuals, inbred, produced progeny which hatched
in May and became adult in July and August in the following ratio :

Long Winged 224 : Short Winged 7 {F„).

These F^ short winged individuals, inbred, produced progeny in Sep-
tember, which, after great mortality, matured the next March and all
were short winaed.

R. K. Nabours

163

Going back to tlie long winged forms from the inbreeding of the
Fi generation short winged, their progeny which hatched in August
matured from December to February :

Long Winged 20 : Short Winged 8 (F^).

These last short winged did not produce progenj' until the following
March and they grew from March to June resulting in:

Long Winged 10 {F,).

The ^1 generati<in long winged gave progeny which hatched in May
and became adult in June and July and were :

Lung Winged 336 : Short Winged 4 (F^).
DIAGRAM II.

/ 33 (/.■,,)

July long toimj

June

April

1910-^

65 (/'V
hmtj wing

10 {F,)
long wing

1909 J.

58 (
Marcli long

Feb.

Jan.

Dee.

21;

Sept. long

Auk.
July
June
April
March

5{F)
short tvi'iig

190S Dec.

7 (/■i)
short imno

hulchcd short wing x short wing

This is nearly the same result as that obtained from the inbreeding
of the Fi generation short winged brothers and sisters. From the
inbreeding of the short winged of this last fraternity, progeny were

164 Inheritance and Evolution in Orthopteni T

hatched in August which matured from December to February and
gave:

Li>ng Winged 15 : Intermediate Winged 2 : Short Winged 41 (F,^.
Now going back to the long winged of generation F., long winged,
their inbreeding gave progeny in July which matured from September
to February :

Long Winged 212 : Intermediate Winged 2 : Short Winged 25 {F-^.
The long winged in this instance were nearly all matured by the end
of September, while the intermediate and short winged required until
February to mature. The short winged ones just enumerated as
maturing in February, produced young in March, which matured in

June :

Long Winged 9 {F,).

The inbreeding from the long winged of F, generation progeny
produced young in October which matured the following March and
April :

Long Winged 58 : Intermediate Winged 3 : Short Winged 68 {F^.
These short winged gave young in April which matured in July :

Long Winged 65 {F^).
Their long winged brothers and sisters, generation F^, also produced
young in April which matured in July :

Long Winged 83 ( F^).

V. An examination of the location, arrangement, and rela-
tions OF the pigmental elements in the colour patterns
of the extracted species and their hybrids.

All the hybrids thus far produced, except some of those in which
texanus {A) is a component, exhibit, on supei-ficial examination as
shown in the photographs, the colour charactei-, or part of it, of each
of the components. In some cases, as leuconotus-leucothorax {BC),
leuconotus-melanothorax (BG), and leucothorax-melanothorax (CG), on
superficial examination the elements of the parents appear to be present
in apparently equal pioportions, while in others, as ieuconotus nigro-
notatus (BI), leuconotus-luteonotatus (BH), leucothorax-nigronotatus
{CI), leucothorax-luteonotatus (CH), leuconotus-rufrolineatus (BF),
leuconotus-luteolineatus {BE), leucothorax-iiifiolineatus (CF), and

R. K. Nabours

165

leucothorax-luteoiineatus (GE), the characters of the leiiconotus {B)
and leucothorax (C) parents appear to obscure or replace the brown
or mottled brown parts of the characters of nigronotatus (/), luteo-
notatus {H), rufrolineatus {F), and hiteolineatus {E), respectively, and
allow only their more brilliant parts (mahogany brown, yellowish red,
and dense black) to appear (Plate VI). It was first thought that these
more brilliant parts (mahogany brown, yellowish red, or dense black)
were separated, in the patterns of the hybrids, from the grays or mottled
brown parts which accompanied them in the patterns of the parent
species. In order to test this matter more definitely than a superficial
examination of the patterns allows, sections have been made through
the pronota of several of the hybrids and their parent species and these

Fig. 3.

1(3() Tnlieritcmce and Evolution in Orilioptero 1

examined microscopically. The examination of the section through the
pronotum of nigronotatus (/), about midway between the deep black
spot and the posterior end of the pronotum, reveals a deeply pigmented
hypodermis with the cuticle somewhat brown in places (Fig. 3, 7).
The examination of a sfection of leuconotus {B) from approximately the
same location as the one taken from nigronotatus (/) (Fig. 3, E) shows
the hypodermis to be practically without pigment and the cuticle
colourless. A section from approximately the .same part of the prono-
tum of leuconotus-nigronotatus (^7) (Fig. 3, BI) reveals a nearly clear
cuticle with about one-half as much hypodernial pigment as is found
in nigronotatus (/). The situation regarding pigmentation in the pro-
notum of texanus and the hybrid from it and leuconotus {B) appears
to be exactly the same as that for nigronotatus (7), leuconotus (5)
and their hybrid leuconotus-nigronotatus {Bl). In the figure (Fig. 3)
texanus is placed opposite nigronotatus, and the leuconotus-texanus
hybrid is placed opposite the leuconotus-nigronotatus {BI) hybrid,
although the drawings were made from the specimens first described.

This study reveals the fact that the character of nigi-onotatus (7) is
as much present in this posterior part of the pronotum of the hybrid
leuconotus-nigronotatTjs {BI), as the more advantageously displayed
leuconotus {B), though the latter when the hybrid is scrutinized super-
ficially, is the only one apparent. The same pi'oportions and relations
in the pigmentation of texanus and the heterozygote, leuconotus-
texanus, are shown. The preliminary examination of the pronota of
some of the other hybrids and their parent forms reveals a similar
situation. The evidence indicates that the 23eculiar pigmental elements
of each of the patterns of the pronota of the parents are present in the
pronotum of their hybrid in about ecpial proportions.

VI. Discussion.

(1) r//e inheritance of the colour patterns. The inheritance be-
haviour throughout the experiments, with five exceptions (leuconotus-
luteolineatus-nigronotatus {BEI ) Table III (e)), and four others now
being bred and to bo described later, fulfils very nearly the Mendelian
expectations. Among more than 5000 recorded individuals resulting
from the crossing of species, inbreeding and crossing of hybrids, and
the crossing of hybrids with species, and more than 2000 recorded
progeny from the inbreeding of species, only the five unexpected

E. K. Nabocrs 167

individuals appeared, aud the expectations in regard to the propor-
tionate numbers have been fairly realized.

The Mendelian assumption that hybrids do not produce gametes
representing themselves, but give gametes of the species from which
they themselves were formed and that these gametes are produced
alternatively in about equal proportions, accounts, with the five
exceptions, for all the results which have come from my breeding
experiments with the grouse locusts.

In Exps. I (F,-), II (&) and (c). III (a), (6), (c), (d), and V (a), (b).
the ancestry of the parents used was known for one or more generations,
and their resulting progeny, in the matter of patterns completely (with
the five exceptions in thousands), and in the matter of proportionate
numbers, fairly, approximated to the expectations of alternative in-
heritance. As the results from known parents are closely approximate
to expectation, it seems reasonable to expect the equally regular and
similar results from parents whose ancestry was not known at first
to lead to the identification of the constitution of the parents them-
selves. It has been by this method that the constitutions of the
parents from the field and group cultures used in Exps. I, II, and IV
have been determined.

(2) The appearance of lone/ and short winged n ess. A glance at the
behaviour of the wing lengths character shows that the short winged
required the maximum of time to reach maturity after hatching, and
that this great length of time is closely correlated with the time of the
year — the fall and winter months. The long winged individuals on
the other hand required a minimum of time to reach maturity after
hatching, and this minimum time is also closely correlated with the
time of the year — the spring and early summer. The time of the year
during which growth proceeds seems to determine whether it shall
extend over a long period or not. If the time for growth be a long
one the wing lengths are likely to be short; if the time required for
gi'owth be a short one the wing lengths are likely to be long. The
length of the wings of the parents does not condition this character in
the progeny. The progeny of short winged individuals become long
winged if they grow quickly in the spring. The progeny of the long
winged individuals become short winged if the growth take place
slowly during a long time. Long winged individuals may produce
a majority short winged if the growth take jDlace from October to
April, while their brother and sister short winged ones may produce
all long winged, if the growth progress from March to June. Nor does

168 Inheritrnice and Evolution in Orthoj>fera J

tlie phenomenon appear to be due to an inherited seasonal rhythm ; tor
the fourth generation progeny, coming from the short winged genera-
tion III, which had grown from July to February, the time required
for two generations of their brother and sister generation II and III
progeny, behaved exactly as the progeny F^ of the long winged and
short winged which had come from the fourth generation of the same
line.

(3) Equivalence in the hybrids. From the examination of the
pigmental compositions of the colour patterns of the pronota so far
as it has progressed, the conclusion seems to be justified that the
peculiar pigmental elements of each of the patterns of the pronota of
the parents are present in the pattern of the pronotum of their hybrid
in about, if not in exactly, equal proportions. With this knowledge in
mind it does not appear that the terms dominant and recessive are
applicable at all to these grouse locusts ; they appear to be, in respect
to their representation in the composition of their hybrids, perfectly
equivalent, or, to use Davenport's term, equipotent (2). If only the
superficial appearances be taken into consideration, Bateson's terras of
cpistatic, for the colour most apparent, and hypostatic, for the colour
less apparent, may be employed in some instances (1).

The fact that the heterozygote pattern in the end result is so
equivalently made up of the respective patterns of the parent species
seems to warrant the suggestion that the somatic part of the hybrid
zygote (fertilized hybrid ovum) in its somatogenesis may be in some
way alternative, giving the character of the one, and then the character
of the other, parent to the resulting soma of the hybrid ; just as the
gametal part of this same liybrid zygote in its gametogenesis is usually
most certainly alternative, giving a gamete for the one, and then a
gamete for the other parent.

(4) The " Genotype Conception." These forms approximately, if
not completely, fulfil the requirements of the description of biotypes
by Johannsen. The evidence points to the fact that in none of the
inheritance behaviour observed is there any transmission of the qualities
of the parent to the offspi-ing(5). (There have been five exceptions
noted.) The regular 1:2:1 ratio result of the inbreeding of hybrids,
the 1 : 1 ratio I'esult of the crossing of hybrids with their parent types,
and the 1:1:1:1 I'atio result of the interbreeding of hybrids indicate
that the qualities of the parents, as well as the qualities of the progeny,
are determined by the nature of the germinal material, and that the

R. K. Nabours 16J»

germinal material of each species is pure and inviolate from generation
to generation, whatever the combinations that are made with them.
The fact that two germ plasms come together to make a heterozygote
does not alter this situation, because, although combined in fertilization
into a harmoniously acting zygotic system, they immediately separate
in gametogenesis as though they had not been mixed at all but had
been held together only. The resulting soma (the pattern only con-
sidered here) indicates that each of the gametes gives the soma
characters of its own kind, and that these two sets of characters, from
the two parental sources brought together in fertilization, are in a sense
dove-tailed one into the other to make the individual heterozygotic
combination.

VII. Conclusion.

The inheritance behaviour of the colour patterns in these ortho-
pterouH insects shows clearly the Mendelian type of inheritance, and the
essential result of these experiments has been the extension of this
principle to a considerable number of types of a phylogenetically low
group of ametabolous insects.

All the hybrid patterns, except a few which have not been
adequately examined, show plainly in their visible somatic constitution
all the parts which can be distinguished in the somatic make-up of
each of their parent patterns. No character of one parent species is
ever replaced in the F^ hybrid by any character of the other parent.
All the characters of each parent are represented in the F^ hybrid.
It follows, then, that these grasshoppers do not exhibit characters,
which by crossing can be replaced by other different characters; the
whole pattern appears to be the only unit.

Dimorphism and polymorphism in the length of the wings and pronota
are not inheritable, but are somatic, due to variable incident conditions
under which the individuals grow. The conditions causing slow growth,
extended over several months, produce a preponderance of short winged
individuals. These conditions obtain in the fall and winter, and may
be a matter largely of the lack of sunshine. The conditions causing
quick growth, extended over a shorter time, produce a preponderance
of long winged individuals. These conditions obtain in the spring and
early summer and may be largely a matter of an abundance of sunshine.

I desire to express my thanks to Prof W. L. Tower for the use of
equipment and for much valuable time given in consultation during
the progress of the work and the preparation of the MSS. The late

170 Inheritance and Evolution in OrtJio/dcrd I

Prof. C. O. Whitman first suggested the problem and introduced me
to Dr J. L. Hancock who has given vahiable aid in the Taxonomy and
Natural History of the material. To these friends I am grateful for
invaluable assistance and encouragement.

EXPLANATION OF PLATE VI.

Most of the photographs are of females, the patterns of both males and females being
the same. The difference in sizes is not significant, being due to the scale of the
photographs.

The top row represents tlie true breeding forms. The middle and lower rows repre-
sent eighteen of the heterozygous forms derived from crossing the pure strains. C'G, C/,
FVL, and EH are modified photographs, the patterns of the rest being untouched or only
tinted.

BIBLIOGRAPHY.

1. 1909. Bateson, W. '' Mendel's rrinciplcs of Heredity." Camliridgc Uni-
versity Press.

2. 1907. D.^VENPOHT, C. B. " Heredity and Mendel's Law." Proceediiujs of
the Washington Academy of Sciences, Vol. IX. p. 179.

3. 1910. DoNCASTER, L. " Heredity in the Light of Recent Research." Cam-
bridge University Press.

4. 1902. Hancock, J. L. "The Tcttigidae of North America." Chicago.

5. 1911. JoHANNSEN, W. " The Genotype Conception of Hcrcdity." American
Naturalist, Vol. XLV. No. 531.

6. 1907. LuTZ, Frank E. "The Variation and Correlation of the Taxonomic
Characters of Gryllns." Carnegie Inst. Pub. 101.

7- 1910. Montgomery, T. IL " Arc Particular Chromosomes Sex Deter-
minants?" Biological Bulletin, XIX. No. I, pp. 1 — 17.

8. 1910. Tower, W. L. "The Determination of Dominance and the Modi-
fication of Behaviour in Alternative (Mendelian) Inheritance by Conditions
Surrounding or Incident ujiou the Germ Cells at Fertilization." Biological
Bulletin, xvill. pp. 285 — 337.

9. 1893. Whitman, C. 0. "Tlie Inadequacy of the Cell Theory of Develop-
ment." Woods Hole Biological Lectures.

JOURNAL OF GENETICS, VOL. IIL NO. 3

Pure Strains

Paiatettix texanus
Hano.

Hybrid

P. leuconotus,
n. sp.

P. leucotliorax,
□ . sp.

P. iJUuctofemorata,
n. sp.

AB

BC

BG

Bl

Hybrids

CF

CE

CH

CI

PLATE VI

\

P. lufrolineatus,
n. sp.

/

P. melanothorax,
n. sp.

P. luteonotatus,
u. sp.

P. uigi'onotatus,
n. sj).

BE

CG

Dl

\. y

EH

El

EF

A PRELIMINARY NOTE ON THE GENETICS OF

FRAG ARIA.

By C. W. RICHARDSON.

The work described in this note was carried on at the John Innes
Horticultural Institution, and I am deeply indebted for the facilities so
kindly afforded me. This work is as yet in a preliminary stage, but
enough progress has perhaps been made to justify an inteiim report.

§ I. Experiments with F. vesca.

(a) Alpines {F. vesca semperflorens).

Alpines are generally said to belong to the vesca species, yet they
differ from the English vesca in two or three minor habits of growth
and in particular in the important habit of continuous flowering.
Amongst the numerous varieties of Alpines is one, F. de Gaillon, intro-
duced by Labaute in,lf^ll, which never produces stolons (runnei-s). At
present my work on the nature of this runnerless condition is scarcely
advanced enough to publish. But runner x runnerless always gives
runner-producing plants in F^, and runner and runnerless in F.., the
runner being a mai'ked dominant. The DR plants produce fewer
runners in their second season than is the case with normal plants.

Fruit Colour.

In 1910 I crossed runnerless White (W) with runner-producing
Red {R).

TABLE I.

1910

Parents W ^ R

1

1911

Fi

1

R

1

i

1912

F.,

70 Red

1

i

20

White

\ (no intermediates)

3 to

1 Expectation

67-

oR,

22-5 W

172 A PreUminari/ Note on tlie (fenetics o/Fragaria

{[>) F. vesca mouophylla.

Duchesne's father sowed elatior (Hautbois) seed in 1760, and in
1761, amongst the plants that came up, sowed vesca seeds. The plants
were neglected till 1763, when Duchesne fils went through them and
found one plant with single leaves (Fig. 1). This plant he said brrd
true whether from seed or stolons.

Fig. 1. Fig. 2. Fig. 3.

In 1910 I received .some plants of »no»o^j/<?/iia character from France;
some of these in spring and late autunui produce a few bifoliate and
trifoliate leaves. In 1912 I selfed one plant, apparently a pure iiiuno-
phi/lla, and obtained 15 plants, up to the present, always true, and
10 plants giving occasional bifoliate or trifoliate le;wes (Figs. 2 and 3).

In 1911 I cros.sed vesca, trifoliate, normal (N ) with monophylla
abniirnial (^4).

TABLE II.

1911 Parents N x A

I
I'.ll'i Fi N

I

I 1

i I

V.IU F.. 177 N 73 A (Plate VII, tig. 1)

Before leaving the subject of monophyllas I would call attention to
a curious plant (Plate VII, fig. 2). Before I had any monophyllas I selfed
a. plant that had "gone wild " on the edge of a gi-ass-covered bank invad-
ing a bed of R(jyal Sovereigns ; from this I obtained a large family not
unlike a family I obtained at the .same date from a selfed Royal Sovereign.
From one plant of this family which I selfed I obtained a small family
all more or le.ss abnormal. The plant depicted is probably the most

JOURNAL OF GENETICS, VOL. IN. NO. 3

PLATE VII

Fig. 1.

Fig. --i.

C. W. Richardson 173

abnormal. As may be seen it produces leaves like those of monophylla,
others bifoliate and trifoliate in no set order. This plant, though in
its third season, has as yet produced no Howers.

§ II. Garden Hybrids.

Something of the origin of the modern Garden Strawberry is known,
but its whole history is not. It springs fr(3m an old form of Garden
Strawberry, the results of crosses between vescas, Alpines and Hautbois.
This older form was again crossed with F. virginiana, introduced in 1629,
and F. chiloensis, introduced to Marseilles in 1712 and to England in
1727 by Philip Miller. These crosses were again crossed with F. anan-
assa (F. grandiflora^), introduced fi'om Holland during the eighteenth
century. The origin of this plant is unknown. It was said to have
been brought fi-om Surinam, where to-day there are said to be no straw-
berries. It is also said to have been a variety of virginiana brought
from Carolina. It may be a chinensis cross, as Holland received many
plants from China during the eighteenth century. Whatever its origin
all our best garden varieties of to-day are descended from Fr. ananassa
crosses. I have selfed 8 varieties of garden fruit, producing over 1000
plants. Not one resembles a vesca, or an Alpine, but many show distinct
traces of chiloensis, more of virginiana and not a few of chinensis. The
leaf-character of Hautbois occasionally appears in those of French origin,
and I have met with it in the offspring of " Latest of All " — a variety
derived by Laxton from the French Helena Gloede as one parent.

Perpetuals.

There is no precise record of the parentage of the first perpetuals.
It is generally stated that they were crosses of garden varieties and
Alpines — the perpetual habit (i.e. the habit of flowering and fruiting
more than once in a season) coming from the Alpines. I have crossed
Alpines with garden varieties, but have invariably found the resulting
plants produce very poorly developed flowers, which have no pollen, and,
when crossed back with their original parents, produce very few seeds,

1 Miller's Figures of Plants, Vol. ii. 1760. — "Some Persons have affirmed it" (Fr.
ananasse) "was brought from Louisiana; others, that it came from Virginia; but I
received some Plants of this Kind from a curious Gentleman of Amsterdam, who assured
me they were brought from Surinam."

Cf. Duchesne, Histoire Natnrelle des Fraisiers, p. 190, Paris, 1766. Also J. Gay,
Annales des Sciences Naturelles, viii. 1857, p. 204. Also Knight, Tr. Hort. Soc. iii. p. 207.
None of these accept the Surinam theory of the ' ' curious Gentleman."

•lourn. of Gen. iii 12

174 A Preliminarji Note on the Genetics o/Fragaria

The plants I have of F^x P are not <ild enmigh to flower, but the
results should be interesting. This at least tends to discourage the
view that Alpines cannot be crossed with garden varieties, though, at
the same time, it does not prove that our perpetuals of to-day sprang
originally from the cross Alpine and garden variety.

The first perpetual widely known was " Gloede's seedling," Ananas
Perpetual, no great success. Then came Mabille (Limoges) with I'ln-
epuisable, also no great success. Then came Roi Henri, St Joseph and
the perpetuals of to-day the work of Abbe Thivolet.

In the endeavour to segregate the perpetual character I selfed, in
1911, St Antoine de Padoue. In 1911 I counted 93 Perpetuals (P)
and 35 non-Perpetuals {NP), 2 doubtfuls. In 1912 I tested these 130
plants again removing all flowers to prevent early fruiting and obtained
the result in September and October

108 P 22 NP

Expectation (3 to 1) 97 -.5 P S2-5 NP.

In 1910 I selfed "Laxton's Perpetual."

In May 1912 I removed all flowers fi'om the resulting plants and
obtained in October,

69 P 11 A^P 2 doubtful

Expectation (3 to 1) QVo P 20-5 NP.

In 1910 I crossed the garden variety "Bedford Champion" (non-
perpetual) with " Laxton's Perpetual."

In May 1912 I removed all flowers from the resulting plants and
obtained in October,

24 P 53 NP.

The simplest expectation would be equality, from which the de-
parture is considerable. Later results suggest that \'arious complications
have to be considered which, as yet, I cannot interpret. Subsequent
experiments have resulted as follows : —

In 1911 "St Antoine de Padoue" was again .selfed, and gave in
October 1913—

64 P 6 A^P 1 doubtful, also 16 dead plants.

Perpetuals appear to resist drought better than non-perpetuals,
accordingly it is quite possible many dead plants were non-perpetual.
Another point, worth noting, is that this year (1913), after dry weather
during July, August and September, many garden varieties flowered in
October which would not have done so under normal conditions.

C W. Richardson 175

I find Fo'fi of my first St Antoine family fi-equently come true tu the
perpetual habit. But no clear case of a non-perpetual F^ breeding true
has appeared — the nearest gives

■20 NP 2 P 2 possibly NP.

For reasons already stated the two P's may be NP's.

F^'s of " Bedford Champion " x " Laxton's Perpetual " have resulted
up to the present in

F, (Parent HP)

8 NP 6 P

F, (Parent P)

14 NP 5 P.

With the pure perpetuals, now obtained, more satisfactory results
are to be looked for, but the evidence to hand points to the perpetual
habit depending upon more than one factor.

§ HI. Sex.

In crossing Fragaria the question of sex inheritance is of the gi-eatest
importance. It is rash to come to the conclusion that any cross, made
with plants of one sex, which produces sterile flowers, is bound to
produce sterile flowers when made with similar plants of a different sex.
Ivnowledge of the heredity of sex, if it could be obtained, would be of
extreme use to the producer of new garden varieties, for the best fruit
comes from the plants that have both the male and female parts of the
flower well developed'.

Various sexual arrangements are found amongst strawberry floweis.

1. Females with the male organs undeveloped.

2. Females with most of the female organs atrophied or hyper-
trophied and inefficient and no male organs developed.

3. True hermaphrodites with both male and female organs de-
veloped.

4. Males with the female organs undeveloped.

5. Males with the female organs only developed in a few flowers
(generally the first flowers produced in each truss).

6. Flowers with neither male nor female organs developed, or with
female organs h3^ertrophied.

' Keens remarked on Hautbois with a dioecious arrangement. Tr. Hort. Soc. ii.
p. S93.

12— -2

176 .4 Preliminary Note on the Genetics q/Fragaria

My first virginiana plants, obtained from Kew, were females with
rudiments of stamens but no pollen. Accordingly I obtained plants ft'om
America, which were said to be males. Any that flowered the first
year they arrived were apparently males. In the following year (191.3)
I found these new plants consisted of males, hermaphrodites and females.
Some apparent hermaphrodites are really males, but as these cannot be
distinguished from functional hermaphrodites by simple inspection,
I have as yet been unable to make the crosses Male x Female,
Male X Hermaphrodite and Female x Hermaphrodite, and to self Her-
maphrodite ; but at the earliest possible date I intend to make these
crosses, as my plants are now apparently classified.

The following crosses made in 1911 are of interest (purely female
virginiana used throughout).

Parents virginiana and chiloensis (I think) pure male.

F, 16 ? \-2 ^ 6 ? .

Three plants did not flower.
Parents virginiana x chiloensis hicida, pure male.

F, 49 ? 27 </ 16 ? .

Eleven plants did not flower.
Parents virginiana x grandiflora, hermaphrodite.

20 ? 0 ^ 14 9.

Forty-two plants did not flower.
Parents virginiana x vesca, hermaphrodite.

^1. A very few fertile females, a very few fertile hermaphrodites,
sterile females, some apparently males and a large number of plants
that did not flower. (The plants are rather too young to base results
on but show three sex-types.)

A cross of nearly pure male chiloensis hermaphrodite x grandiflora,
hermaphrodite (used as male parent) gave a minority of hermaphrodite,
a majority of males, and no females.

That the crosses female virginiana x grandiflora hermaphrodite
and nearly male chiloensis x the same grandiflora pollen produced no
males in the first case and no females in the second is certainly curious.

It is worth remarking that one or two female flowers on a plant
may set a seed or two and all the other flowers be quite sterile, or
again on another j^lant of the same parentage one female flower may
set seed freely and all the other flowers be sterile.

C. W. Richardson 177

Fig. 4 represents an effort of a sterile female to prcjduce a fruit
without setting a single seed, though it has been thoroughly polli-
nated'. The actual cross here is vesca x Laxton's Bedford Champion.

if-

Fig. i.

A few plants of this cross have set a few seeds when pollinated with
either parent ; of the two The Champion seems the more successful
sire ; at present these cross-backs are too young to draw conclusions
from.

The cross pink-flowering vesca x tuhite vesca produced pink F^'h. I
have F„ plants which should flower next spring. The cross double x
single flowering vesca produced plants with single flowers and occasion-
ally flowers with a few extra petals in F^. My FJs should flower next
season. The thick rubbery leaves of chiloensis have appeared in all F-^'s
I have obtained from this species crossed with others, though the leaves
are larger than those of the iiovvsibX chiloensis ; some i'Vs of my chiloensis
crosses should be old enough next year to show to some extent how far
leaf characters can be segregated. Some multifoliate plants when selfed
have given a majority of multifoliate plants. My actual numbers were
too small to base conclusions on. On the whole, I expect, judging by
results I have already obtained, that most characters possessed by
Fragaria are capable of segregation, but at the same time some linked
characters may present difficulties which it may take years to solve.

For plants of F. virginiana fi'om a vnld source I am indebted to
Dr W. A. Orton of the Department of Agriculture, Washington, and
Professor Czapek of Prag very kindly sent plants of F. elatior which I
am also using.

' Knight mentions a seedling Hautbois x Alpine in which the fruit developed
though no seed was formed. Tr. Hort. Soc. v. 294.

PARTHENOGENETIC AND SEXUAL REPKUDUC-
TIUN IN SIMOCEPHALUS VETULUS AND
OTHER CLADOCERA.

By W. E. agar.

University of Glasgow.

In 1879 Weismann t'onnulated the theory that in the Cladocera "der
Eintritt der Geschlechtsperiode wird tiberhaupt nicht durch direct
wirkende aussere Momente veranlasst, sondern lediglich durch innere
in der Natur des Organismus begrtindete." There can be no doubt that
this total denial of the influence of environment can no longer be main-
tained. It is as certain as experiment can ever make it, that under
some conditions parthenogenetic reproduction will proceed longer than
under others, and that to some degree the appearance of sexual forms
can be controlled by controlling the environment. The latest and
probably the most conclusive evidence of this is given by Grosvenor
and Smith for Moinu. It is true that many experimenters have failed
to find any evidence of the direct effect of environment, but this
negative evidence cannot be put against the quite satisfactory positive
results of other workers. On the other hand, even those who admit the
influence of environment do not give up the idea of an underlying
" cycle " as well (Issakowitsch, Papanicolau, Woltereck, R. Hertwig).

The object of this note is to present some new evidence and point
out some general indications that the change from parthenogenetic to
sexual reproduction, and the degeneration which so often accompanies it
under experimental conditions, is determined by environment only, and
that the number of preceding parthenogenetic generations or the lapse
of time since the last sexual act are not, as such, relevant to the matter
at all.

My own experiments have dealt chiefly with Simocephalus vetulus,
and were not undertaken in relation to the reproductive cycle problem,

180 Reproduction in Siinocei)halus vetulus

but in connection with xariout;; problems of heredity. As, howcvor, the
experiments have been extensive, and as one line in particular was bred
for 46 successive generations, they are able incidentally to afford very
instructive information upon the sexuality question.

It would be useless to review completely the bulky literature dealing
with rejjroduction in Cladocera, and for this I refer the reader to the
discussions and literature lists in the works of Papanicolau, Woltereck
and Hertwig. It is well, however, to mention a few researches dealing
specially with S. vetulus.

This species was one of those dealt with by Weisraann in 1879, and
on which his theory quoted above was based.

Issakowitsch (1908), who experimented with this among other species,
comes to the conclusion that it has an inherent reproductive cycle, the
duration of which is not, however, independent of external conditions.
For the " cycle " is due to an increasing " depression " of the germ cells
(in the nucleo-cytoplasm relation sense) which in the early stages can
be alleviated by various external factors, but at last gets to a stage when
nothing but conjugation will save it from death.

Kuttner (1909) supported Weisinann's extreme view. S. vetulus was
one of the species experimented with.

We must consider Papanicolau 's experiment (1911) rather more
closely as it was carried out in great detail and thoroughness, and as
the conclusions have been accepted by R. Hertwig and used by him
in support of his theory of " depression " periods leading to sexual
conjugation.

Papanicolau's main experiment dealt with a line bred from an
ex-ephippio female at room temperature, and gave the following chief
results.

1. The line bred parthenogenetically for twenty generations, sexual
and degenerate forms getting more and more numerous, the twentieth
generation consisting entirely of males and degenerated animals, and
parthenogenesis then ceased altogether.

2. Sexual individuals began to appear in late broods of early
generations (e.g. in the tenth brood produced by generation 2, in the
eighth brood produced by generation 3) and appeared in earlier and
earlier broods in later generations, till they occurred in the second
brood of generation 17 and first of generation 20.

3. A gi'eat many weak and degenerate forms appeared, often
abnormal and dying before birth. The appearance of these degenerate
forms accompanied that of the sexual ones, and the same phenomenon

W. E. Agar 181

of their curlier and earlier apjjearance in succeeding generations was
observed.

4. There was a gradual decrease in the number and an increase in
the size of the eggs fi-om generation to generation, and also, as regards
size, from brood to brood of the same generation.

He therefore concludes that there is an inherent "sexual cycle" of
the orthodox kind in S. vetulus, beginnuig with vigorous parthenogenetic
reproduction which gradually gives place to increasing sexuality and
degeneracy, ending in total cessation of asexual reproduction. He is,
however, prepared to admit that external conditions influence the
duration of the cycle to a certain extent.

The line of S. vetulus whose sexual condition I wish to describe
originated fi-om a parthenogenetic female taken from one of the
laboratory tanks in September, 1911. I have of course no knowledge
of the number of generations which intervened between it and the
last ephippial egg.

Some of the descendants of this individual formed the material on
which I did some experiments recently published on the transmission
of environmental effects, and also an experiment on parthenogenetic
inheritance, not yet published. The method of breeding employed for
the great majority of the individuals was that which I described as
" normal " or " control " in my former papei'. They were bred in
cylindrical glass tubes 10 x 3 cms., which when corked (as they always
were) contained about 50 c.c. of water and 15 c.c. of air. With rare
exceptions one specimen only was kept in each tube, and when it had
produced a brood, this was removed within 36 hours. The culture
medium was the water from a tank in which were living a number of
Lepidosiren paradoxa, fed daily with Anodonta. The water in this
tank was changed weekly (except for that held in a thick layer of mud
at the bottom of the tank) and was thick with Infusoria and organic
detritus. A jar of water was taken from this tank on alternate days,
strained through linen to remove Rotifers, Lynceids, etc. which swarmed
in it, and used to renew the water every alternate day in all the breeding
tubes. Each Simocephalus was picked out of its tube mth a pipette,
the tube emptied and refilled -ndth the new water and the specimen
returned.

The sexual conditions of this line are shown in Table I.

The sex of a new-born S. vetulus is very difficult to determine, and
I did not attempt to do so, as the experiment was not directed towards
this question. The sex of every individual which was kept to maturity

182 Reproduction in Simocephalus vetiilus

TABLE I.

N ainber of Brouilti dud liitlividuals lexled fur /Sexuality in each

Generation.

Gene- 1st 2nd 3rcl 4th 5tli 6th 7th 8th 9th 10th Uth

ration liroods liroods Broods Broods Broods Broods Broods Broods Broods Broods Brood

■1 1—1 1—2 X X —_._____

3 X 3-4 1—1 1—4 _ — — _— — —

4 4—13 X X X— — — — — — —

5 7—21 5—13 2—13 1—6 1—1 x x — - — —

6 "S— 11 4—5 4—10 2—2 2—2 x 1-1 _ — — —

7 *11— 25 5—10 2—2 X x— — — — — —

6' 11—11 X 2—2 4—10 X — — — _ — _

9 *8— 15 1—9 X X X XX— — — —

10 10—26 X 1—2 X __— — ___

11 9—20 9—17 5—11 1—2 X x — — — — —

13 18—36 X X X X— — — — — —

13 24—82 3—12 1—3 1—2 x —— — — — —

2i 38—72 1—3 3—9 1—2 x — _ — — — _

15 31—62 X .^ X X— — — — — —

16 28—28 X — — — — — — — — —

17 26—26 — — — — — — —— — —

in 24—48 X X X X X X X 8—8 — —

19 39—44 X X X X X X X 1—2 1—2 —

30 X 6 — 29 X X xxxxxx 1 — 3

31 11 — 37 X xxxxxxx 1 — 2 —

33 6—9 X XXX 2—4 — — — — —

34 2—4 — — — — — —_—__

35 2—3 — —_— — — — —__

as X 3—12 — — — _—_„ — _

37 9—31 — — — — — — — — — —

3b 13—17 — — — — — — — — — —

39 6—7 — — — — — — — —__

30 2—4 B— 3 — — — _— — — — _

31 3—3 — — — _ — — — —.__
33 3—10 — —-— — — — —__

33 5—7 — — — _- — — — —___

34 5—9 — — — — — — — —__

35 3—4 — — — — — —____

36 1—6 — — — — — — — —___

37 3—7 — — — — — — — — — —

3S 3—4 — — — — — — — — — —

39 4—4 — — — — — — ____.

40 3—3 — — — — — — — —__

41 3—3 — —.—_— — ___
43 3—3 — —_— — — — ____

43 3—3 — — — — — _ — —__

44 3—3 — — — — — __ — —__

45 3—3 — — - _ — — — —__
,^6'x — — — — — — — —__

In each column the number of broods (families) tested is given first, and then the

number of individuals from these broods which were allowed to come to maturity
(see text).

X , Brood recorded, but not tested for sexuality.

* Males occurred in these broods (see text).

W. E. Agar 183

was however recorded, and these form the material available for the
table. One example of how the table is to be read will explain it at
once. Take generation 10' for example. In this generation ten first
broods were tested for sex — i.e. the first broods from ten individuals of
the preceding generation. By "tested for sex" I mean that one or more
of the young were brought to maturity and the sex recorded. In this
case altogether 26 individuals were taken fnjm the ten broods. In the
same generation no individuals belonging to second broods were kept
to maturity, but one third brood was tested by the isolati(3n of two
members of it. As the members of a brood in Swiocephalus (as in
Daphnia) generally all, or mainly all, belong to the same sex, it is
important to state how many broods were tested, and not merely the
number of individuals without reference to the number of families from
which they were taken.

The re.sult of the experiment was as follows : — No sexual (ephippial)
females appeared during the whole of the experiment. Males appeared
in three generations only, where the asterisks are placed in the table.
In generation 6 there was one male from one of the first broods, in
generation 7 two of the first broods furnished between them five males,
and in generation 9 there were three males from one of the first broods.
All the other broods and all the other generations afforded partheiio-
genetic females only.

Thus so far from a progressive increase in sexuality, the very few
sexual fonns that did appear were in the early generations of the
experiment.

These 46 generations were passed through in twenty-one months,
at room temperature. The average duration of each generation
would however have been far greater at the mean annual temperature
in their natural ponds. Papanicolau estimates the maximum number
of parthenogenetic generations in a natural " cycle " at twelve, and this
agi'ees roughly with my estimate of the greatest number that could be
produced in a year under the climatic conditions of Glasgow. Even
during the time in which the line was bred in my experiment, partheno-
genesis lasted therefore for nearly four times as many generations as
may be expected in nature (though of course the natural " cycle " is by
no means a rigid one), and at the end of the experiment no trace of loss
of vigour was detectable.

According to Papanicolau, not only should sexual forms appear with

1 The original parthenogenetic female from which the Une was started being counted
as generation 1.

184 Reproduction, in Simocephalus vetiilus

increasing frequency in later generations, but also in later bronds of
earlier generations. Now all the four broods which gave males in lay
experiment were first broods. Yet broods of a much higher order were
tested and furnished parthenogenetic females only.

Also according to Papanicolau, a potent factor in producing sexual
forms is not only the order of the brood, but the oixler of the brood to
which the parent belonged. The greater part, but far from all, of each
generation in my experiment was descended from first broods in the pre-
ceding generation. This however was also the case in Papanicolau's main
experiment (Tafel I), in which the line ended altogether in sexual and
degenerate forms in the twentieth generation. A direct test of the
influence of the order of the brood was also made. It will be seen that
in generation 22 two sixth broods were tested for sex. Now the parents
of these sixth broods were members of the tenth brood of a specimen
of generation 21 which was itself a member of the eleventh brood of an
individual of generation 20, itself descended from the tenth brood of
generation 19, the parent of which was a member of the ninth brood of
generation 18. This is a far severer test than any which Papanicolau
describes, and yet all were parthenogenetic females. (Breeding from
late broods like this makes each generation fall far behind, in point of
time, the generations bred chiefly fi'om first broods. The sixth broods
of generation 22 were contemporary with the first broods of generation
36 of the main part of the line.)

The third of our list of Papanicolau's conclusions has been no more
lay experience in this experiment than were his other.s. He found
degenerate broods extremely common, increasing in frequency pari
passu with the sexual forms. I have had very few of these degenerates,
and they certainly showed no tendency to increase as the line grew
older.

As mentioned above, the gTeat majority of the individuals of this
line were bred under the conditions which I have for convenience
termed " normal " as distinct from the intentionally abnormal conditions
under which a part of the line was bred. The " normal " and abnormal
environments resulted in similar sexual conditions however. The
abnormal environments included feeding with a protophyte culture
which produced certain specific abnormal characters (described in my
paper of 1913), cultivation in chemical solutions, and at high and low
temperatures. In the case of the high temperature, part of the line
was subjected to the abnormal condition for four or five successive
generations. Naturally abnormalities and a higher death rate were

W. E. Agar 185

produced by these conditions, but in the main part of the line, under
the normal conditions, " degenerates " have been extremely rare.

On two occasions however there was an epidemic of deaths among
young individuals and embryos in the part of the line under normal
conditions. The first of these occun-ed in generation 4, the second in
generation 30. The second one was very instructive as at that time
I had under identical conditions parallel cultures of two other lines
started from specimens obtained from different localities to that from
which the parent of the principal line was obtained. The two lines in
question were started from parthenogenetic females from Cambridge
and Beith respectively, and generation 13 of the Cambridge line and
generation 14 of the Beith line were contemporary with generation 30
of the main line, and all of them showed simultaneously the same
"degeneration." In the main line four females laid altogether 19 eggs,
from which only two living young were obtained. In the Cambridge
line, six females laid together 31 eggs, and only 12 living young were
born, while from the Beith line no living young were produced from
29 eggs laid by six females. These figures all refer to the first broods
of the parents. All three lines had completely recovered by the second
broods of the same generation, the second batches of eggs laid by all
these sixteen females developing perfectly normally into normal healthy
young. Moreover, except for occasional cases here and there, no more
" degenerate forms " appeared in any of the three lines. In fact in the
main line (Table I), in the last eleven generations not a single one of
the new-born young isolated failed to arrive at maturity, and to produce
perfectly normal young. Clearly therefore the temporary and simul-
taneous disturbance in the three lines was due to some temporary
unfavourable factor in the environment, although there was no reason
to suppose that the water in the Lepidosiren tank varied in any
way from the normal at that time ; while all the other factors of the
environment were, so far as could be ascertained, unaltered.

The last of the visible accompaniments of degeneracy observed by
Papanicolau which we need consider is the diminution in number and
the increase in size of the parthenogenetic eggs as the line gi-ew older.
These are really largely the same phenomenon, there being a strong
organic correlation between the number and size of the eggs, the larger
the number the smaller being therr size. This is the ease, that is to
say, if we judge the size of the eggs by that of the new-bom young
hatched from them. I have found it very difficult to get a satisfactory
series of measurements of the eggs owing to their rapid increase in size

186

RejJrodKction in Simocephalus vetulus

as they develop. The yonng however do not grow between ecdyses to
any extent detectable by the micrometer. An actual determination of
the coefficient of correlation between size of young and number of eggs
for S. vetulus gave — "74 + '04. The value for S. exspinosus, worked out
in larger numbers, gave as an average — •48. The values were computed
for groups for individuals belonging to the same generation and all
giving birth to theii- young nearly at the same time, and refer to first
broods only.

As these experiments were primarily directed to the inheritance of
certain dimensions of the young, I have a large number of measure-
ments available. Table II gives the mean length of the new-born young

TABLE II.

Mean Length of the New-Born Yowkj {in first Brooda only).
Unit of Measurement = 0'018 mm.

Oene-
mtioii

Mean Length

of New-Hom

Young

Xumber

of
Broods

Number

of

Individuals

Gene-
ration

Mean Lengtli

of New-Born

Young

Number

of
Broods

Number

of
Individuals

3

43-717

3

23

2o

43-667

4

30

4

44-448

4

29

26

44-654

3

13

5

43-4.5.5

2

11

27

43-954

12

76

6

43-4.50

4

20

28

45-277

23

195

7

4.5-G43

2

7

29

44-(;-29

11

58

.1

43-911

0

28

30

45-000

1

1

9

43-333

C

24

31

45-133

3

15

10

43-27n

5

29

32

46.500

2

4

11

43-191

9

44

33

43-714

3

28

12

41-628

7

39

34

42-071

3

14

13

43-9.5-2

15

104

35

44-050

3

10

14

43-616

62

468

36

47-083

1

6

15

41-736

64

639

37

43-547

3

21

16

42-194

62

474

38

45-864

3

11

17

44-896

28

208

39

not measured

18

44-279

26

231

40

not measured

19

43-384

43

241

41

43-115

3

13

20

43-263

42

302

42

44-017

3

30

21

41-.536

29

223

43

44-692

3

26

22

41-310

37

179

44

43-667

3

12

23

44-000

8

47

45

43-278

3

27

24

44-350

4

20

46

44-848

3

23

Note. The column "Number of Individuals" gives the numbei; of youns measured
in each generation, not the total number of voung born. When any brood contained
less than ten individuals all of them were measured, but when there were more than ten,
only the first ten, taken at random. Thus the mean number of young per brood cannot
be obtained from this table by dividing the number of individuals in each generation
by the number of broods.

W. E. Agar 187

(i.e. in their first instar), for the last 44 generations, the first two not
being available for this purpose. Only Ji^st broods arc dealt with in the
table, and those jDortions of the line living under the abnormal conditions
described above are omitted, as these conditions directly affected the
size of the new-born young.

It will be seen that the size fiuctuates greatly, but there is no real
tendency to a progressive increase. If the forty-four generations are
divided into four gi'oiips of eleven each, we find :

Mean length for generations 3—18 43'637

14—24 43-017

25—35 44-423

36—46 44-457

It will be seen that the second set of eleven generations average
smaller than the first. It is true that the last period gives the highest
average, thanks to generation 36 which was represented by only a single
brood of exceptionally large individuals. The mean of the last six
generations however is 43-936, almost exactly equal to the total mean
(43-889) and so is the mean for the first six, 44-104. There is thus
no justification for supposing that the size of the young is increasing,
although by taking twenty successive generations (Papanicolau's number)
a good case might be made out for either increasing or decreasing size
according to the portion of the table included by them.

I can fully confirm Papanicolau's statement that the size of the egg
(or new-born young) increases from brood to brood — at any rate up to
about the fifth. Exact measurements of this increase were given in my
former jDaper. This increase however is in no sense due to a cyclical or
cumulative action of the increasing lapse of time since the last sexual
act. It is a phenomenon which begins afresh in each generation, and
seems to be a common occurrence throughout the animal kingdom. The
increasing size of the eggs of successive ovipositions has been shown in
detail by Halban for Salamandra maculosa and S. atra, and also for
Anura and Ghelonia. It is the experience of every poultry keeper that
the young hens lay smaller eggs than the older ones, and it is well
known to be the case with man that the size of the new-bom infant
tends to increase in successive parturitions.

Whilst the line described above was in progress I also bred numerous
others, the most important being two started respectively from a par-
thenogenetic female kindly sent me from Cambridge by Mr F. A. Potts
and from one from Beith, 16 miles from Glasgow, kindly sent to me by

188 Reproduction in Simocei)halus vetulus

Mr John Ritchie. From the Beith specimen 17 generations were taken
and fi'om the Cambridge specimen 16 generations, giving with the
original females 18 in the Beith and 17 in the Cambridge lines. They
were kept under exactly the same conditions as the main line, and in
neither of them were any sexual forms found at all, nor any appearance
of " degeneracy," though as described above a very large proportion of
the eggs of the first broods of the contemporary thirteenth generation
of the Cambridge, and fourteenth of the Beith lines fixiled to develop.

It again happens that if the period during which these lines were
bred is divided into two portions, the mean size tif the yi>ung in the
earlier generations is less than in the later ones. Combining the con-
temporary generations of the two lines together we find that the mean
length of the first brood young of the first period (seven generations
measured) is 45"256, and of the second period (also seven generations
measured) is 45'904. A closer analysis shows however that it would be
again quite unjustifiable to conclude that a progi-essive increase was
taking place. If we an-ange the generations in order of the magnitude
of the new-boni young, we find the order, beginning with the smallest,
is (generations 1 and 9 not being available)

8, 5, 16, 2, 11, 17, 4, 3, 10, 14, 6, 12, 15, 7.

In a fairly extensive experience of breeding Cladocera I have of
ctjurae often experienced the phenomena of bursts of sexuality and
of " degeneracy " — or better, of high mortality. I have had both these
phenomena appearing in D. pulex, and the sexuality in iS. exspinosiis
kept under the conditions described as normal above, and also in
S. vetulus kept under other conditions.

It is obvious therefore fi'om this and from the work of others that
the same sj)ecies may under certain conditions exhibit an increasing
tendency to sexuality and degeneration ending in total cessation of
parthenogenetic reproduction, and under others may continue the
asexual reproduction apparently indefinitely. My main line was to all
appearance as vigorous at the 46th generation as at the first, and so
were the Cambridge and Beith lines as long as they were bred — and it
must be remembered that the parthenogenetic females which originated
the three lines may themselves have had a long parthenogenetic
ancestry. There is no justification for the supposition that if the
experiment had been continued " long enough " the lines would have
at last been unable to maintain themselves without sexual repro-
duction.

W. E. Agar 189

Three conditions of the environment seem to be specially significant
in interpreting the ahnost uniform lack of sexuality, and the total lack
of any tendency to degeneration observed in these three lines under long
continued parthenogenesis.

Fii'stly, with the few exceptions of those individuals fed with the
protophyte culture, they were all fed with the same food, and this food
supply, though it may have fluctuated from day to day, probably did
not do so over long periods, as the water in the Lepidosiren tank (which
had been established more than three years before the experiment began)
was changed weekly. Hence any cyclical change in the food supply was
probably a weekly one, and there was little chance of a progressive
deterioration in the culture medium lasting over weeks and months
which might have caused sexuality and degeneration to set in.

Secondly, practically every individual was isolated in a separate
tube within 48 hours of birth. The only excep)tions were so rare as to
be negligible, and moreover in only one case where more than one
individual was kept in the same tube was such a sj)ecimen used as the
ancestor of any considerable number of generations. In the light of
Grosvenor and Smith's results as to increased sexuality in overcrowded
animals, this is a significant point.

Thirdly, the water was changed regularly in all the tubes every
second day — with again an insignificant number of exceptions.

General considerations.

It is obvious that it is no longer necessary in the present state of
our knowledge to discuss the extreme Weismannian hypothesis that the
parthenogenetic or sexual mode of reproduction is determined entirely
by internal changes which aie an integi-al part of the physiology of the
animal and independent of environment. The idea of an internal cycle
or rhythm still persists however, and it is undoubtedly the most generally
accepted view to-day among workers on Cladocera that the change from
parthenogenetic to sexual reproduction is determined by such a cycle,
with the limitation that this change can be accelerated or delayed by
particular conditions of the environment. This is expressed by Hertwig
in the following sentence (p. 29): " Fortgesetzte Parthenogenese fiihre
schon ah solche' in der Beschaffenheit der Tiere zu Veranderungen,
welche die Entwickelung der Geschlechtsgeneration veranlassen. Diese
Umformung der Zellen konne durch einen entgegengesetzt wirkenden
Faktor, wie die Warme, vielleicht dauernd zuriickgedriingt werden."

1 My italics.
Journ. of Gen. iii 1^

100 Reprofhtction hi Sinioccphalus vctulus

Amongst other upholders of the view of an inherent cyclical change
which can be influenced to a greater or less extent by environment are
Issakowitsch, Papanicolau, Woltereck. The kind of cycle maintained by
Woltereck is however quite different from that supposed by Hertwig. The
last-named worker correlates it with his nucleo-cytoplasm relation theory
and considers that the lajjse of parthonogenetic generati(jns and of time
leads to a state of depression in the germ cells that eventually under
normal conditions ends in sexual conjugation or death. Woltereck justly
points out that the theory would have to be strained to breaking point
to cover his own experiments with Hyalodaphnia, etc. For instance,
Woltereck bred a line of H. cucuUata parthenogenetically for four years,
during which time it fluctuated between pure parthenogenesis and pro-
nounced sexuality. Hence we should have to suppose that the line
recovered from pronounced sexual depression periods without sexual
conjugation.

Woltereck 's own view is that there are two antagonistic substances
in the egg, the predominance of one resulting in parthenogenetic
individuals, and of the other in sexual forms. These substances are
supposed to wax and wane alternately and autonomously. At such
times (labile periods) as they are nearly equally balanced, external
conditions are able to turn the scale one way or the other, and thus at
times environment influences sex. " Mann kann deshalb die innere
zyMische Periodizitdt der Valenz (of these two substances) mit Recht
als das Kernproblem der Cladocerenfortpflanzung bezeichnen" (p. 123).

Although Woltereck's cycle is fiir more compatible with the observed
facts than is the theory of a depression caused by continued partheno-
genesis as such, nevertheless there does not seem to be sufficient evidence
for the existence of any sort of cycle at all, while there is strong evidence
against it.

The evidence for the existence of such cycles consists mainly of :

1. Experiments where the conditions were supposed to be constant,
such as (a) Papanicolau 's work with Siiiiocephalus and Moina, where an
increasing sexuality and degeneration are observed, and (6) Woltereck's
with Hyalodaphnia, where the line fluctuated between pure partheno-
genesis and a high degi'ee of .sexuality.

2. Experiments where abnormal conditions of various kinds are
found to have a determining influence in some cases and not in others.
When such conditions are effective it is supposed that it is because the
cycle has reached the point where the tendencies to parthenogenesis and
sexuality are nearly balanced. When they are ineffective, it is ascribed

W. E. ArxAR 191

to the fact that the cycle is too near one end or the ot.her for the opposite
condition to be evoked.

Now it is quite possible to account for these observations without
invoking an internal cycle. Firstly, it is extraordinarily difficult to keep
all the conditions of the environment constant, and especially the great
difficulty of keeping food cultures constant for long periods imposes upon
the experimenter, who assumes that physiological changes in his animals
were not correlated with changes in the environment, the obligation of
stating very fully what precautions and tests he took to ensure that the
conditions really were constant.

Secondly, even where the conditions are constant throughout the
whole experiment, an increased tendency to sexuality in, say, the
twentieth generation as compared with the tenth may be due to
the fact that the line has been living for a longer period in an
unsuitable environment — perhaps, for example, one deficient in some
essential constituent or " vitamine." As both Papanicolau and
Woltereck point out, not only the number of generations but also
the length of time during which a line has been subjected to the
experimental conditions is of importance in determining sexuality, for
sexual forms appeared in late broods of early generations and early
broods of late generations. In my experience with S. vetulus an
individual of any generation produces its fourth brood about the same
time as the members of its first broods produce their first brood of off-
spring. That is, the fourth broods of the reth generation are contempoi-ary
with the first broods of the ?i + lth generation. If this held true for
Papanicolau's strain of S. vetulus, it is easy to calculate from his Tafel I
that sexual and degenerate forms appearing in the later broods of the
earlier generations actually appeared earlier, in point of time, than those
of later generations.

I have described experiments (1913) where the effects of a peculiar
environment produced on a given generation of 8. vetulus were still
detectable in their great-grandchildi-en, and Woltereck has produced
evidence to show that an environment acting on an individual may
determine the sexuality of its gi-andchildren. It is plain, therefore,
both that environmental effects may persist for some generations after
they were produced — in other words that it may act in cumulative
fashion — and also that the length of time and not only the number of
generations during which the line has been in the conditions of the
experiment, is of importance in determining sexuality. Taking this in
conjunction with the fact that under some environmental conditions no

13—2

192 Be2))'odi(c(io)i in Simocephalus vetulus

tendency to sexuality or degeneration appears even after an enormous
number of parthenogenetic generations, it seems necessary, on the
principle of accepting the simplest hypothesis which will fit the facts,
to conclude that the sexual cycle (obligatory parthenogenesis — laliijc
period — obligatory or preponderating sexuality, often accompanied under
experimental conditions by " degeneration ") is, when present, due
entirely to the cumulative effect of an unfavourable environment or to
an actual though often unsuspected change from a favourable to an
unfavourable environment.

This view does not, of course, diminish the physiological interest of
the change from the parthenogenetic to the sexual mode of reproduction.
Indeed the tendency to replace asexual by sexual reproduction under
certain conditions (often unfavourabU' ones) is a phenomenon of the
deepest significance, but it seems to be no more due to an inherent life
cycle than the increasing hunger, ending in degeneration and death
unless the conditions are changed, which accompanies the withholding
of food fi-om an organism, is due to an inherent physiological cycle.

The ascertainment of the exact conditions under which asexual is
replaced by sexual reproduction, and the precise advantage conferred
by the latter, is indeed a most important task. The external conditions
under which parthenogenesis will continue indefinitely are certainly
different for different species. They may be rarely realised in nature,
and still moie rarely for any prolonged period, natural conditions being
subject to seasonal and other changes. For many species of the i-elated
Ostracoda however these conditions do seem to obtain in nature. On
the other hand the other extreme is nut impossible, that for some species
in which asexual reproduction is one of the normal modes of reproduc-
tion, there are no conditions under which asexual reproduction can go
on for more than a limited time. In other words, that there is no
environment which does not act in a way prejudicial to parthenogenetic
reproduction. If such species do exist, it might be legitimate to speak
of their sexual cycle, but in that case it would be necessary to remember
that the " cycle " is not due to anything of general import but merely
to the peculiar relations of that particular species to the environment.

It appears that a similar change of view is taking place in regard to
the life cycle of the Infusoria. For many years the orthodox view has
been that originated by Maupas as the result of his classical experi-
ments. According to this view the life history is a cyclical one, the
near descendants of the exconjugate being in a state of immaturity,
which gi'adually yields, as asexual multijDlication proceeds, to puberty,

W. E. Agar 193

ending in senility and to the necessary death, through internal causes,
of the asexual colony. As better and better methods of cultivation have
been evolved the duration of this "cycle" has been gradually lengthened,
and artificial stimuli substituted for the "rejuvenescing" conjugation,
till at last we have Woodruffe, after breeding Paramecium aurelia for
3340 asexual generations without evoking any signs of " degeneration,"
coming to the conclusion " dass das Protoplasma einer einzigen Zelle
unter giinstigen ausseren Umstanden ohne Hilfe von Konjugation oder
einer kiinstlicher Reizung imstande ist, sich unbegrenzt fortzuptlanzen
und zeigt ferner in klarer Weise, dass das Altern und das Befruchtungs-
bediirfiiis nicht Grundeigenschaften der lebendigen Substanz sind "
(p. 36).

Similar conclusions have been reached by Jennings as the result of
extensive experiments on individuals of Paramecium which had conju-
gated, and others which had been prevented from conjugating though
ready to do so.

Conclusion.

The following conclusions, though refen-ing particularly to S. vetulus,
may probably be safely extended to a wide range of the Cladocera.

1. Certain not yet fully elucidated factors in the environment
influence the onset of sexuality.

2. Certain factors likewise bring about " degeneration " or high
rate of mortality.

3. Certain factors of the environment may act cumulatively over
a number of generations.

4. Therefore the increasing sexuality and " degeneration " (or high
mortalit}') observed inidei' certain supposedly constant experimental
conditions receive a ready explanation in the supposition that the
environment is one favourable to the development of these phenomena.

5. This explanation is made much more probable when we find
that under other experimental conditions there is no tendency to
increasing sexuality or degeneration.

6. Many species exhibit the phenomenon of specially labile periods,
when sexuality is easily influenced by certain factors of the environment.
This labile condition is usually asci'ibed to the fact that the line is in
about the middle of the reproductive cycle, the diminishing tendency
to parthenogenesis being about equally balanced by the increasing
tendency to sexual reproduction. Such a balanced condition must
however be passed through equally whether the tendency to sexuality

194 Reproduction in Simocephalus vetulus

is being increased by the progress of the " cycle " or by the cumulative
effect of an unfavourable environment. Hence the existence of labile
periods is as readily explained on the one hypothesis as on the other.

7. There is no justification for retaining the hypothesis of an
inherent reproductive cycle — that is to say, the hypothesis that the
number of generations or lapse of time since the last fertilised egg
influences, as such, the production of sexual or degenerate forms. For
the production of these forms is under certain conditions not influenced
even by the lapse of an enormous number of parthenogenetic generations,
while their production certainly is influenced by environment in other
cases. The residuum of cases being equally well explicable on either
hypothesis (cycle or environment) it is most reasonable to suppose that
the factor that was effective in the one case (environment) was the one
that was effective in the other, and conversely, that the ineffective factor
of the one case ("reproductive cycle") was ineffective in the other.

LITERATURE CITED.

Agar, W. E. " Traiisinission of Environmental Eftccts from Parent to Ofispring

in Simocephalus vetulus." Phil. Trans. Roy. Soc. London, Series B. 203,

1913.
Grosvenor, G. H., and Geoffrey Smith. "Tlie Life-cycle of Moina rectirostris."

Quart. Journ. Micr. Sci. Vol. LVIII. 1913.
Halban, Josef. " Die Grossenzunahme dei- Eier uml Neugeborenen mit dem

fortsclireitenden Alter der Mutter." Archie fur Entwickiungsmech. Vol. xxix.

1909.
Hertwig, R. "tJberden derzeitigen Stand des Sexualitlits|ir()l>lem.s nebst eigenen

Unter»\ichungen." Biol. Centralbl. Vol. sxxii. 1912.
IssAK0WlT.sc'H, A. " Es be.steht eine zyklische Fort[)flanziuig bei den Cladoceren,

aber nicht im Sinne ^\'eismann's." Biolog. Centralhl. Vol. xxviri. 1908.
Jennings, II. S. "The Effect of Conjugation in Paramecium." Journ. E.cp. ZooL

Vol. XIV. 1913.
Kdttner, O. " Unter.suclumgen liber Fortpflanzung.sverhaltnisse und Vorerbung

bei Cladoceren." Internal. Revue der gesamnit. Hydrohiol. u. Hr/drographie,

Vol. II. 1909.
Papanicolau, G. "Expcrimentelle Untersuchungen iiljer die Fortpflanzungsver-

haltui.sse bei Daphniden." Biol. CcntndhL Vol. xxx. 1910.
Weismann, a. "Beitrage zur Naturgeschichte der Daphnoiden." Zeit'tchr. fiir wiss.

Zoolog. Vol. xxxiii. 1879.
WoLTERECK, R." " Vcriinderung der Sexualitiit bei Daphniden." Internat. Rev.

der gesamint. Hydrohiol. %i. Hydrographie, Vol. w. 1911.
WooDRUPPE, L. L. "Droitausend und dreihundert Generationen von Paramecium

ohne Konjugation oder kiin.stliclie Reizung." Biolog. Centralhl. Vol. xxxill.

1913.

ON THE APPEARANCE OF STERILE "DWARFS"
IN HUMULUS LUPULUS L.

By E. S. salmon, F.L.S.,
8.E. Ar/ric. College, Wye, Kent.

In a large number of " crosses " which I have made during the past
seven years between different cultivated female varieties of hops and
various individual male hops', some of the resulting seedlings are
remarkably distinct in character from either parent. The distinguish-
ing features of these seedlings are (1) their total, or almost total,
inability to climb; and (2) their complete sterility, no flowers being
produced.

With reference to the first character, we find that in these abnormal
seedlings the strong relatively thick climbing stems (" bines ") which in
the normal male or female hop-plant arise annually from the perennial
rootstock are replaced by a number (often a very large number) of
weak, thin, sometimes almost thread-like stems, of limited growth,
which are either totally unable to climb, or climb weakly a short
distance. In many of these abnormal seedlings — which may con-
veniently be termed " dwarfs " — all the shoots either grow prostrate
on the surface of the ground or form an erect bush-like growth about
1 foot high ; in other cases the longest shoots if provided with a piece

' Prof. J. Percival {Agric. Botany (1902), p. 335) writes: " ...the male [hop] is alwaj's
practically a wild form, for on account of their being of no use to tlie grower, males have
never been subject to special selection and improvement. It is somewhat curious that
although female seedlings show considerable variation, we have never Been any morpho-
logical differences among males, no matter what their origin, except in cue or two solitary
instances where the ' bines ' were a paler colour than usual. " This statement is somewhat
misleading, since we find in the forms, or varieties, of the male hop quite as much varia-
tion in such characters as the colour of the stem and petioles, length of the lateral
branches, and in other vegetative characters as in the female hop-plant.

190 Sterile " Dvarf^" in Hiinmlus LupuUis L.

of cocoa-nut string (such as is used in hop-gardens for " training" hops)
and "trained" round it from time to time through the summer, will
climb in rather a feeble manner and reach a height of 3 to 5 feet. No
better climbing habit is shown if the shoots are allowed to gi-ow up
finely-branched pea-sticks. In the majority of cases a "dwarf" pro-
duces a large number of thin shoots, which radiating from the rhizome
run prostrate on the ground and produce a " mat "-like growth covering
a square yard or more (see PI. VIII, fig. 1); in rarer cases a few of the
shoots if carefully " trained " raise themselves by climbing round each
other and the cocoa-nut string, either to a height of 5 feet, as shown
in PL VIII, fig. 2, or more usually to a height not exceeding 2 feet
(PI. IX, fig. 3). The leaves on the "dwarf" are much smaller than
those on the normal hop-plant, and are less divided, being often unlobed,
and never more than 3-lobed ; the diameter of the stem at its base does
not exceed ^ in., and is often less than ^ in., whereas in the normal
plant it is from | in. to ^ in. The stem of the " dwarf" possesses the
hooked hairs found in the normal plant. So far as has been observed,
the root-system of the 'dwarfs" is characterised by none of the roots
running horizontally; further, no underground stems gi-owing hori-
zontally (" runners ") have been seen in the dwarfs.

The distinctive characteii sties of the "dwarf" are usually evident
soon after germination. A normal seedling produces in a few weeks
a stem with a strongly developed climbing habit, which, in the case of
a vigorous plant, will reach at the end of the first year's growth, to a
height of .5, or even 6, feet. In cases where " dwarfness " is most
marked, the seedling plant produces in its first year one or more shoots
of very limited growth, which never climb and which bear abnormally
small and often curled leaves ; at the end of the season's growth the
whole plant may be only 1 inch high. In other cases, where the
" dwarfness " is not so extreme, a number of shoots are produced, which
may reach to a length of 6 to 9 inches ; none of the stems, however, are
able to climb. In the second year from germination, the normal seedling,
whether male or female, produces a fairly stout stem, which climbs to
10 feet or more, and produces flowers. A normal seedling developing
the fresh shoots in the spring of its second year is shown in PI. IX, fig. 4.
In the case of " dwarfs," the one-year-plant if vigorous produces the next
spring a number of shoots, with thin .sometimes almost thread-like
stems, which never climb, but run prostrate over the gi'ound and attain
a length of 2 to 3 feet. No flowers are ever produced (as noted below).
PI. IX, fig. 5 shows a one-year-old " dwarf" .seedling, starting the season's

E. S. Salmon 197

growth. In the cases where the "dwarfness" is most pronounced', the
plant remains for the second year and longer extremely stunted, pro-
ducing a number of shoots only a few inches high with very small and
often curled or distorted leaves. PI. IX, fig. 6 shows such a dwarf, at
the end of the second year's growth. Such plants apparently — where no
sjDecial care is given to their cultivation — die after a few years. The
larger " dwarfs," however, may show a vigorous growth in the third and
succeeding years, and may produce a veiy large number of trailing
stems (frequently over fifty) which radiate in all directions to a distance
of 3 or 4 feet. The most vigorous "dwarfs," however, some of which
are now 7 years old, have never produced a single normal climbing
stem, nor reached a height (or length) of more than 5 feet, — whereas
the stems of all the other normal seedlings of the same " crosses " can
climb to 20 feet or so.

" Dwarfs," like the normal plants, are liable to be attacked, and
much injured by "mould" {Sphaerotheca Humuli) and "green fly"
(Phorodon Humuli), and to be killed by "eelworm" {Heterodera
schachtii).

The second characteristic of " dwarfs " is their absolute sterility, no
flowers or rudiments of sexual organs having been produced in any case.
Over 200 " dwarfs," arising from various " crosses," have been kept under
observation. " Dwarfs " have occuiTed among the normal seedlings in
the majority of the " crosses " I have made ; e.g. in 25 " crosses " in which
the following female (English) varieties of hop were used ; — Cobb's
Golding, Colgate, Fuggles, Early Bird, Early White, Canterbury White-
bine. The male hops used were different individuals of the English
forms of the male hop, and in one case a male hop obtained fi'om
Oregon, U.S.A., which differs in leaf- and other characters, from the
English forms.

As some of the " dwarfs " are now 7 years old, and as the normal
seedling produces flowers in the second year after germination, it appears
safe to conclude that the present kind of dwarfness, unlike that recorded
in Humulus japonicus by Figdor^, is associated with absolute sterility.

With regard to the proportion of dwarfs that may occur in the
Fi generation, the following facts are available.

' "Dwarfuess" is most pronounced in those "crosses" in which the male hop is a
form from Oregon, U.S.A., and the female a variety cultivated in England.

2 W. Figdor, Uebergangsbildungen von Pollen- zu Fruehtblattern bei Humulus japoni-
cus Sieb. et Zucc. and deren Ursachen, in Sitzuvgsber. d. hniserl. Akad. d. Wissensch.
Wien, Bd. cxx. Abt. 1 (1911).

198 Sterile "Dwarfs" in Humulus Lupiilus L.

Ill a "cross" (Ref. lui. 5/()7) made in 1907 the f'eiiiak' hop was the
variety known as Fuggles', and the male hop (Ref. no. M8) one of the
English forms with a red-bine. 51 hops containing 256 seeds were
obtained from the pollinated flowers; while the 44 "control" hops
contained no seedsl 87 seedlings were raised, of which 52 j^roved to
be climbers and fertile', and 35 proved to bo " dwarfs " and sterile.
Prof Bateson has pointed out to me that here the ratio of "dwarfs"
to " climbers " approximates to 7:9.

In a second cross (Ref. no. 4/07) made at the same time, between
the same female variety (Fuggles) — though different individuals'* were
used fi'om those in the first cross — and a certain male hop (Ref. no. G 27),
one of the English forms with a green-bine, 99 hojjs containing 285 seeds
were obtained from the pollinated flowers, and 28 hops, all without
seeds, from the " control " branches. 67 seedlings were raised, of which
66 plants proved to be climbers and feitile", and 1 plant a dwarf and
sterile. If we assume that this single dwarf was due to the fertilisation
of a flower by a strange poUen-gi'ain during the time the bag was
removed'', and if we also assume that the female plant in these two
" crosses " was identical in character*, then it appears that the latent
characters of dvvarfness and sterility are carried by some English forms
<:>f the male hop and not by others. However this may be, it is quite
certain that the proportion of "dwarfs" to climbers (if dwarfs really
occurred at all) in this second "cross" was altogether different from
that obtaining in the first " cross."

In a third "cross" (Ref no. 1/09) made in 1909, the female hop was
the German variety " Stirn " and the male hop one of the English forms
with a bine striped with red and gi-een (Ref no. Z 12). The pollinated

' Prof. Pereival {Jourii. Royal Agric. Soc. England, lxii. p. 87 (1901)), writing of the
origin of this variety, says, " The original plant was a casual seedling which appeared in
the flower-garden of Mr George Stace, of Horsmonden, Kent. The seed from whicli the
plant arose was shaken out along with crumbs from the hop-picliing dinner-basket used by
Mrs Stace, the seedliug beiug noticed about the 3'ear 18()1. The ' sets' were afterwards
introduced to the public by Mr Richard Fuggle of Brenchley, about the year 187-5.'"

- The controls, it should be noted, were imperfect in that the bags covering them were
not removed at the time when this was done to the branches that were pollinated.

■' 31 were ? , 4 were <? , and 17 were killed by eelworm before they flowered.

* The commercial cultivation of the hop being entirely vegetative, by means of sets or
cuts — the thickened basal portion of the stem — it might be assumed that all individuals of
any variety are identical ; it is very probable, however, that the stocks of some varieties
have not been kept true.

■' 57 were ? , 7 were i , 1 was monoecious, and 1 was killed by eelworm before it
flowered.

'■ See footnote '2 above.

E. S. Salmon 199

flowers gave 15 hops with 261 seeds, and the "control" branches 58 hops
with no seeds. 120 normal seedlings were raised, all climbers and
fertile^ — no "dwarf" appearing among them.

In a fourth cross (Ref no. 14/09) the English female variety Canter-
bury Whitebine was pollinated from a male hop obtained from Oregon,
U.S.A., which jJossesses vegetative characters distinct from all the
English forms of H. Lupulus. 110 hops, containing 899 seeds, were
obtained from the pcjllinated flowers, and 85 hops, containing 2 seeds,
from the " control " branches. 109 seedlings were raised, of which 79
were climbers and fertile, and 30 " dwarfs " and sterile. In this " cross,"
however, many of the " dwarfs " were extremely stunted, and were weakly
from the first, and died in the first or second year after germination
before a count was made. It is certain, therefore, that the proportion
of " dwarfs " to climbers in this " cross " is considerably higher than
30 to 79, which are the numbers of plants surviving in the fourth year
after germination.

The only apparent reference to " dwarfness " in H. Lupulus which I
have been able to find is in a Bulletin- by Dr W. W. Stockberger and
J. Thompson, where a bare mention is made of the occurrence of " hills
with dwarfed vines " in a Californian hop-field.

Figdor (I.e.) has recorded the occurrence among seedlings of the
annual species Humulus japonicus of some individuals which showed
a dwarfed habit. H. japonicus usually attains a height from the
ground of 1"5 to 2m.; the tallest of the "dwarf" seedlings measured
only 041 m. and the shortest O'll m.

Upon such dwarf plants which seemed to be otherwise males were
developed flowers hermaphrodite in various degTees (for details see
original paper), and in a few cases female flowers also were formed.
Figdor sought to explain the production of these dwarfs as follows : " Der
Nanismus der einzelnen Individuen wird durch die gleichzeitige Ein-
wirkung einer bestimmten chemischen Lichtintensitat bei verhaltnis-
massig niedriger Temperatur und ebensolchem Feuchtigkeitsgehalte
der Atmosphare in Verbindung mit Nahrungsmangel hervorgerufen."

The " dwarfs " of H. Lupulus described above differ in being
absolutely sterile, and whatever maj' be the true explanation of the
appearance of " dwarfs " in H. japonicus, it is certain that " dwarfness "
in H. Lupulus cannot be attributed to the influence of outside factors,

1 108 were ? , 11 were s , and 1 was monoecious.

■^ Some Conditions influencing the Yield of Hops (U.S. Department of Agriculture,
Bureau of Plant IntUistry, Circular No. 56 (1910)).

200 Sterile " Divarfs " in Humulus Lupulus L.

since the "dwarfs" were treated as regards cultivation etc. from the
time of germination onwards in exactly the same way as the seedlings
which developed into normal plants.

EXPLANATION OF PLATES.

PLATE VIII.

Fig. 1. Photograph taken in the Experimental Hop-garden at Wye College, Kent, showing
the climbing bines of normal seedlings of H. Lupulus, and, to the right, two "dwarf
seedlings which have produced a large number of very thin stems all totally unable to
climb. These "dwarfs" are 7 years old.

Fig. 2. Photograph as in Fig. 1 ; in the background can be seen the climbing bines
of normal seedlings, in the foreground a "dwarf" seedling, the most vigorous shoots
of which have climbed to a height of 5 feet. The leaves are cordate and simple, and
may be compared with those of the normal climbing plants shown in Fig. 1.

PLATE IX.

Fig. 3. Photograph of a "dwarf" seedling; the most vigorous shoots have climbed 2 feet.
The leaves are simple and cordate, and the stems thread-like. This "dwarf" is
7 years old.

Fig. 4. A normal one-year-old seedling, starting the season's growth.

Fig. 5. A "dwarf" one-year-old seedling, starting the season's growth.

Fig. G. A "dwarf " seedling at the end of its second year's growth. Natural size.
(This was one of many similar .seedlings obtained in the "cross" Canterbury White-
bine X male hop from Oregon, U.S.A.)

JOURNAL OF GENETICS, VOL. 111. NO. 3

V "Tff., ».-

Fig. 1.

PLATE VI

JOURNAL OF GENETICS, VOL. IN, NO. 3

PLATE IX

■V. i ■'. . ■--'St' ^■'iv

Fig. 3.

Fifj. 4

Fig. 5.

Fig. 6.

NOTE UN THE UFFSPEING OF A DWARF
BEARING STRAIN OF GUINEA PIGS.

By I. B. J. SOLLAS,

Newtiham College, Guinbridge.

The breeding experiments on a dwarf bearing strain of Guinea pigs
described in the Repo)tt> to the Evolution Committee of the Royal Society
(Sollas 1909) have been continued. The results up to the present are
shown in the following tables :

(1) Complete list of all tlm dwarf -contaiuiiuj families^.

Offspring

Female

Male

Female

Male

(1)

286

287

2

3

(2)

400

401

5

14

(3)

409

i'oi

2

—

W

413

410

—

4

(5)

415

416

4

8

(6)

495

497

10

8

(7)

.571

416

—

2

(8)

571

401

3

2

(9)

571

312

7

5

(10)

571

497

1

5

(11)

571

741

4

4

(12)

572

369

1

—

(13)

572

416

10

11

(li)

572

497

4

3

(15)

586

416

—

2

(16)

586

550

3

3

(17)

644

312

1

2

(18)

644

416

1

—

(19)

644

741

6

2

(20)

718

r

3

1

(21)

718

550

1

1

(22)

718

497

1

1

(23)

718

312

1

4

(24)

718

741

2

3

(25)

A'

X

—

—

(26)

735

416

2

—

(27)

735

497

1

1

(28)

735

741

5

1

Totals

Female

Dwarf Normal, sexes

80

90

24

Male unrecorded

1 —

9 18

1 —

3 —

4 —
6

40

Normal 192 ; Dwarf 64. Females 104 ; Males 130.
' Errors in the previous table have been corrected.

22

202 Offspriufi nf a Dirarf Hearinr/ Strain of Guinea Pifjs

The proportion of iioniuil offsju-ing to dwarfs is 3:1. That the
mimbers in the total are in exactly this proportiijn is of course merely
a coincidence.

The 2)repondei'ance of male dwarfs and of the total niuiiber of males
is very considerably diminishc<l since the last report.

If we con.sider separately the offspring in i'ainilies in which the
dwarfs were all male, in those in which the dwarfs were all female ;ind
in those in which the dwarfs were of both sexes, we finil that the excess
of males is confined to the first set. of families, i.e. those in which the
dwarfs were all male. In this total the males are more than three
times as numerous as the females. In the earlier experiments most
of the families were of this kind; the diminution in the excess of males

(2) FdiiiUifS contain.) tiAj divatfo of both t<e.i:es.

offspring

Parents Normal Dwarf

Female

Male

Female

Male

Female

Male

6)

495

497

10

8

2

()

(9)

571

312

7

5

1

3

(13)

572

416

10

11

o

3

(14)

572

497

4

3

1

2

(19)

644

741

6

2

3

1

(24)

718

741

2

3

1

2

Totals ... 39? 32 <r 11? 17 <f

Total 50 ? : 49 <? .

(•J) Fainilivti coiitainiiuj bcsitle)! iioriital uffspriiKj mnU dwarj/i only.

Offspring

r

'iirents

Normal

Dwarf

Female

Male

Female

Male

Male

(1)

286

287

2

3

1

(2)

400

401

5

14

'.)

(3)

409

401

2

1

1

(1)

413

411)

—

4

3

(5)

415

116

4

8

4

(B)

571

401

o

2

2

(22)

71S

497

1

1

1

Totals ... 17? 33 <? 21 <?

Total 54 [f : 17 ? , a fairly close approximation to a 3 : 1 ratio.

I. B. J. 80LLAS 203

is due to the fact that" our present total is made up largely of the
offspring of families of the other two kinds.

In families in which dwarfs of both sexes have already occurred the
numbers of the sexes are equal. In families in which up to the present
no male dwarfs have been born there is an excess of females.

In making out the tables of families containing dwarfs of both sexes
we are clearly safe in including all families; but in both the other cases
we cannot be certain that any of the families, if they had increased,
would have remained in the class in which we have placed them.
Nevertheless the facts may have some interest.

Thus in the families containing dwarfs of the male sex only, the
proportion of the male to the female offspring approximates to 3:1.
It is possible that the 18 normal offspring of unrecorded sex in family 2
(see Table 1) would have altered this proportion; but even if we make
the unlikely supposition that they were all female we should still have
a large excess of males. In family 22 each parent has, when paired
with other mates, produced female dwarfs ; of the other parents 416
and 571 have produced female dwarfs when paired with other mates.
Possibly further breeding would show that some of the families in this
table are out of place.

(■4) Faini/ie-s cuiiiitiuiuij feinah ilwdrfs onbj.

Otfspring

Parents

Normal

Dwarf

^nviiiMl iif

Female

Mali;

Female

Male

Female

unrecordeil sex

(lu)

SS6

416

—

2

1

4

(16)

.■186

550

3

3

2

—

(17)

614

312

1

2

1

—

(21)

718

550

1

1

1

—

(23)

718

312

1

4

2

—

(25)

X

A'

—

—

2

—

(28)

735

741

5

1

4

—

Totals ... 11? 13 c? 13? 4

In this table, the female parents in the matings 15, 16, and 28 have
never yet given birth to any male dwarfs, the males with which they
were mated have had male dwarfs among their offspring. The parents
in the matings 17, 21, and 23 have all, when paired with other mates,
had male dwarfs among their offspring.

Sturtevant (./. Exp. Zool. 12, 1912, p. 513) has published an
interesting attempt to explain these results, as far as they went in

204 OffspriiHj of a Dvarf Bearlmj Strain of Guinea Pigs

the previous report, as a case of parti;il sex linkage. His explanation
accounts for the occurrence of some families containing besides normal
offspring male dwarfs only, others containing besides normal offspring
both male and female dwarfs. According to his scheme there would be
no complete families containing none but female dwarfs. So tar there
may be nothing to conflict with the facts, but as far as I can see, in
both his families the numbers of the sexes would be equal, though he
does not seem to recognise this difficulty himself (.see p. 515).

The experiments will be continued by Miss N. M. and Mr J. B. S.
Haldane.

ON A CASE OF UNILATERAL DEVELOPMENT
OF SECONDARY MALE CHARACTERS IN A
PHEASANT, WITH REMARKS ON THE IN-
FLUENCE OF HORMONES IN THE PRODUC-
TION OF SECONDARY SEX CHARACTERS.

By C. J. BOND, F.R.C.S., Leicester.

Description.

The skin now described is that of the white ringed Formosan
variety {Phasianus Tm^quatm) of the Chinese pheasant.

On the left side (cf. PL X, figs. 1 — 4), it shows the plumage and
spur of the male bird. The red portion of skin round the eye is much
larger on the left than on the right side. The head and neck feathers
are blue green, iridescent on both sides but more so on the left. There
is a semilunar patch of the white collar feathers strictly limited to the
left side of the neck. The wing primaries and coverts are of the female
type though a few male plumage feathers appear in the left wing
coverts. The tail coverts show marked male characters, more especially
on the left side. The breast feathers, which are rufous in colour,
especially on the left side exhibit the black tipping seen in the male
bird. The left leg shows a well developed spur, pointed, and of the size
found in the cock of the second year (cf PL XI, fig. 5). On the right
leg there is no spur but only the usual rudiment or scaur found in the
hen pheasant (PI. XI, fig. 6).

The bones of the left tarsus and phalanges are longer and thicker
than the corresponding bones in the right leg.

The tail feathers (rectrices) are especially interesting. While the
assumption of male plumage is more or less confined to the left side
in the head and neck and trunk and limbs, it is not so limited in the

Journ. of Gen. iii 14

2(l(i Secoudart/ Male Cliai'acteri< in a Plteasant

case of the tail feathers. In the tail (as far as can be ascertained owing
to the unfoi'tunate absence of the two central feathers) it is the outer
half of each feather that has assumed the male type of plumage (PI. XII,
fig. 7). In other words instead of the tail as a whole showing a
division into a male half and a female half, each feather of the tail
shows male colouring and pattern on the outer or less covered half,
and female colouring and pattern on the inner or more covered half

Examination ok the internal organs.

Careful dissection shows the presence of a well developed oviduct
opening into the cloaca in the usual situation on the left side.

In the situation of the ovary (though somewhat nearer to the
middle line) a sex gland was found roughly J inch long by | inch
wide, nodular on the surface and in parts deeply pigmented (PI. XIII,
fig. 10). No trace of a sex gland could be found on the right side.
Microscopical examination of this gland shows actively growing areas
of tubular gland structure in some of which bodies like spermatozoa
can be detected; the acini are lined by columnar epithelium (PI. XIII,
fig. 12). Smaller areas of fibrous tissue with much pigment and degene-
rating cysts are found in another part while a third portion resembles
in structure the cortical part of the suprarenal gland. This sex gland
is in fact an ovci-testis, and from the degenerating pigmented condition
of the ovarian portion (PL XIII, fig. 11), and the actively growing
character of the testicular portion, it seems probable that it functioned
originally as a female gland, and that the ovarian portion sub-
sequently atrophied while the tubular gland or male portion then
became functionally active.

Remarks. The assumption of male secondary characters on one
side of the body only in this case presents considerable difficulty when
we attempt to apply the usual hormonic explanation to the development
of these characters. And yet the facts already established compel us
to accept the hormonic explanation as a partial although not a complete
solution of the problem.

The older experiments on the results of castration in the male
animal and the more recent observations of Marshall, Shattock, and
others on birds and animals conclusively show that with the removal
of all traces of the sex gland in the male the secondary male characters
fail to appear if they have not already begun to grow, or undergo a

C. J. Bond 207

shaqj and peniiaiient arifst of gruwth if they have already begun to
develop bei'ore the castration was performed.

Such are the facts, as experimentally determined, but the nature
of the association, and the way in which the removal of the sex
gland influences the growth of the tissues, are not fully known.

Recent observations suggest that it is not merely the absence of one
sex gland that brings about the development of the opposite secondary
sex characters, but that there must also be the starting into functional
activity of some hitherto dormant elements in the degenerating gland
or in some other situation in order to bring about the evolution of the
corresponding sex characters.

In other words, as Shattock has suggested, the presence of male
sex gland elements is necessary for the development of secondary male
characteristics in every individual whether the female sex gland be
present or not. The essential point is the presence of a stimulating
factor rather than the absence of an inhibiting factor. There are some
facts which support this view thus :

(a) In an increasing number of cases in birds in which atrophy of
the ovary has been found in association with the development of
secondary male characters, careful microscopical examination of the
degenerating female gland reveals the presence of islands of male
elements showing active growth.

PI. XIII, fig. 13 shows such male gland elements in the degenerating
ovary of a hen pheasant in which male plumage and well developed
spurs were both present.

(6) It is known that the gradual destruction of the normal tissue
of both ovaries by a new growth in young women is more often associated
with the growth of hair on the face, alteration in the voice, and the
appearance of other secondary male characters than is even complete
bilateral oophorectomy, the explanation being that the gradual destruc-
tion of the Graafian follicles and interstitial cells by disease acts like
parasitic castration and affords an opportunity for the growth and
functional development of dormant rudiments of male gland cells
present in the degenerating ovary, while in complete bilateral oopho-
rectomy these male elements are removed along with the female gland
and all possibility of their subsequent growth is thereby prevented.
The same seems to be true of birds also. Mr Cooper of Knaresborough
has kindly sent me the photo of a Blue Leghorn Pullet which developed
a well marked comb and other male characters (but no spurs) ; in this
case the ovary was destroyed by a large new growth,

14—2

208 Secondari/ Male Characters in a Pheasant

(c) Although the complete removal of the sex glands at an early
age in the male does no doubt prevent the development of secondary
male characters in most vertebrates, it does not produce the develop-
ment of female secondary characters except in so far as these resemble
those of the young or inunature male.

In the few recorded cases (chiefly among birds) where males have
taken on female plumage, either completely or in part, the change has
accompanied some (at present unknown) natural cause independently
of castration and unassociated in most cases with any atrophic change
in the testes (Hammond Smith, The Field, Feb. 25, 1911). It is
possible therefore that the assumj)tion of female plumage may be
associated with the growth of some dormant ovarian elements in some
abnormal situation. In any case the more frequent occurrence of the
change in the female is of interest when we recall the supposed hetero-
zygous constitution of the female in respect of sex.

Infiuenced by these facts Geoffrey Smith and Mrs Haig Thomas
(Journ. Genetics, June 1913) suggest a different causation for the
secondary sex characters in each case. They suggest that the case
of the female assuming the characters of the male sex is one of
" Correlated Differentiation " and is directly associated with changes
in the corresponding sex gland, while the case of the male assuming
female characters is due to the " Hereditary Transference " of characters
originally associated with one sex to individuals of the opposite sex, and
they instance the hen plumage of the Sebright Bantam Cock as a case
in point.

But even supposing it to be true, as seems likely, that the case
of the female assuming male characters and the case of the male
assuming female characters rest on a different footing as regards
causation, this specimen still falls into a different category be-
cause the assumption of the opposite sex character is limited to
one side of the body in this individual. While it seems clear that
such a localized change of plumage cannot be entirely dependent on
the circulation in the blood of any internal secretion it is also clear that
the explanation of opposite sex characters by Hereditary Transference
is also insutticient, because in this individual bird the sex characters are
different on the two sides of the body. Moreover, not only are the
somatic characters different on the two sides but the sex gland is also
a dual gland composed ol male and female elements. The bird is in
fact a true genetic as well as a somatic Hermaphrodite. It should
I think be placed in a separate class along with those described by

O. J. Bond 209

Brandt {Zeitschrift fur tviss. ZooL, Vol. XLViii, 1889, page 101). Weber
quoted by Biedl mentions the case of a bird (a chaffinch) with right sided
male and left sided female plumage in which a testis was found on the
right side and an ovary on the left side of the body. Virchow (also
quoted by Biedl) records the case of a human pseudo-hermaphrodite, in
whom a left sided anomaly of the sex gland was associated with im-
perfect growth and development on the left side of the body. There
is in the Royal College of Surgeons Museum a butterfly (one of the
Blues) with male wing pattern on one side and female on the other,
but I am not aware that any microscopical examination of the gonads
has been made in this case.

Heiniich Poll ("Zur Lehre von den Sekundaren Sexual Characteren,"
Stzgsher. Ges. Naturf. Freunde zu Berlin, 1909), refers to three cases
of true hermaphroditism associated with a hemilateral distribution of
secondary sex characters in birds. He gives a full and careful descrip-
tion of a bullfinch in which the plumage of the breast was male in
character on the right side and female on the left, and this hemilateral
distribution of secondary sex characters was associated with a double
sex organ, testicular or male on the right side and ovarian or female on
the left side of the body. Poll directs attention to the suggestive
fact that in true hermaphrodite birds with hemilateral secondary sex
characters the male character occurs on the right side of the body,
the side on which the sex gland is also male. He suggests that the
presence of the testis on the right side may be associated with the
atrophy of the right sex gland which normally occurs in birds.

In the case of the Pheasant here described however, the male
characters are present on the left side of the body, while the male and
female elements are gathered together in one single sex gland which lies
on the left side but nearer to the middle line than usual.

Among the Invertebrata, especially the Lepidoptera, abnormal
examples of hermaphroditism occur somewhat frequently. In such
cases the association between the relative position of the internal sex
gland and the distribution of the secondary sex characters is not how-
ever apparently so close as in birds.

Thus while true Hermaphrodites do rarely occur in which male
secondary characters are present with a male sex gland on one side of
the body, and female secondary characters with a female sex gland on the
opposite side, there are still reasons for thinking that the localization of
sex gland and sex character in different halves of the body both have a
common genetic cause, and are not causally related the one to the other.

210 Secondar// Male Characters in a Plieasanl

If this be true of these Hermaphrodites it is also probably true of normal
individuals, all of whom are Hermaphrodites in some degree. In other
words in addition to the sex gland with its Hormonic secretion, we
must also recognise a somatic or peripheral factor in the causation of
secondary sex characters. No amount of Hormone will bring about the
development of secondary male characters in individuals, and in tissues,
in which the rudiments of such characters are absent. On the other
hand sex gland and somatic rudiments being given in heredity, the due
development, and the continued growth of the sex character, depend
on the presence of the sex gland and on the integrity of its internal
secretion.

Without the help of some such somatic, or tissue factor, it is very
difficult to explain these unilateral cases by any Hormonic theory even
if we invoke the aid of the nervous system.

There are some further facts which support this view. Thus, as
Shattock has pointed out, all secondary sex characters are not equally
dependent for their growth and development on the internal secretions
of the corresponding sex glands. See the case of the male Leghorn
chick in the Royal College of Surgeons Museum in which the pre-
cocious development of the testes was associated with a fully formed
male cumb, but in which this testicular activity failed to bring about
the growth of spurs, or sickle tail feathers, another male character.
Comb and spurs and sickle tail feathers stand on a different footing
then in this respect, and it is also known that hens occasionally develop
spurs while continuing to lay fertile eggs. The somatic tissues which
contain the rudiments of the secondary sex characters differ in their
susceptibility to the influence of the corresponding sex gland Hormone,
and it is quite conceivable that in these hemilateral cases tissues which
are genetically homozygous in respect of the male or female character,
tissues which have as it were received a double dose of femaleness or
maleness, are more susceptible to the influence of their corresponding
Horuione than tissues which are heterozygous (nr singly dosed) in
respect of such sex (lualities.

Examination of the tail feathers.

The peculiar arrangement of the pigment pattern in the individual
tail feathers of this bird is of interest in this connection. From a
cursory examination of some skins in the Natural History Museum
S. Kensington, I am inclined to think that hemilaterality of male
plumage pattern in individual tail leathers goes with hemilaterality of

C. J. Bond 211

secondaiy sex characters in the bodj' as a whole, and there can I think
be Uttle doubt that this depends on some important law of ontological
development in birds. In the head and neck, trunk and limbs, the
development of the pigmentary pattern and feather structure seems to
be under the control of separate unilaterally acting factors, whereas
in the tail (at any rate in some birds) one single factor controls
the development of the tail as a whole, and this single factor may
differentiate later into secondary factors which control the development
of each half of each tail feather, as in this case.

There are some interesting facts in neural physiology which bear on
this point. Langley and others have shown that unilateral stimu-
lation of the central nervous system produces unilateral action in the
sympathetic pilo-motor fibres which supply the trunk and limbs, whereas
in the tail the effect of the unilateral stimulation passes over the middle
line and affects the tail as a whole in the cat.

Rorig of Frankfort has recorded cases in which unilateral castration
in the stag has been associated with abnormal gi'owth of the antler on
the opposite side of the body, and other cases uf injury to the fore leg
with arrested gi-owth of the antler on the same side, and cases of injury
to the hind leg with arrested growth of the antler on the opposite side.
There is a specimen in the Royal College of Surgeons Museum which
shows the same thing. The question of the influence of unilateral
castration and unilateral injury on antler gi-owth would well repay
further inquiry. It is possible that secondary sex characters in the
stag are peculiarly sensitive to nervous influences, and it would seem
at any rate that if Rorig's observations are confirmed they cannot be
entirely explained b\' any purely Hormonic theory.

It is interesting to compare this hemilateral feather pattern in the
tail of this hermaphrodite bird with a case described by Shattock and
Seligman in the Transactions of the Path. Soc. LVII, 1906, whei'e they
figure the tail feather of a cock pheasant hatched in 1902 (cf PI. XI,
fig. 14). At Christmas 1903, this bird, while showing the usual adult
male plumage, began to develop female markings at the proximal end of
the central tail feather. This localized assumption of a secondary female
sex character did not come about through the moulting of the male
feather and the growth of a female one, but by the appearance of the.
female j)attern at the base of the male feather. The portion of the
f(?ather so marked was sharply divided transversely from the male portion
and included the whole width and both sides of the feather. During
subsequent moults in 1904-.5 there was ncj reappearance of the female

212 Seconflarji Male Chanieters in a Pheasant

pattern, the bird at that time showing the full adult male plumage and
functioning as a male.

It is I think reasonable to conclude that the gametic factors (what-
ever their nature) in this case controlled the pigmentary pattern of this
tail feather alternately along male and female lines, whereas in my
hermaphrodite specimen they controlled the pigmentary pattern in
opposite halves of each tail feather coincidently.

But there is yet another point of interest about the tail.

In the Royal College of Surgeons Museum the skin and the internal
organs are preserved of a tiue hermaphrodite Leghorn fowl. The micro-
scopic structure of the double sex gland in this case has been cai'efully
worked out by Shattock ; it undoubtedly constitutes an ovo-testis. In this
specimen a well-developed male comb and spurs are present, but there
are no sickle feathers and the tail as a whole (though rather long) is of
the female type. The distribution of the male sex characters is arranged
on an antero-posterior or segmental rather than a bilateral pattern.

There would seem then to be two main t3rpes of sex character
manifestation, as far as the body and limbs are concerned.

Type I. In which a double sex gland is associated with a hemi-
lateral distribution of secondary sex characters, as in my specimen.

Type II. In which a double sex gland is associated with an antero-
posterior or transverse distribution of secondary sex characters, as in
this White Leghorn fowl. These two Types might be di.stinguished as
the Lateral and the Segmental Types.

Although both halves of the tail seem to function in heredity as one
whole (no one has seen a tail with one half male and the other half
female), yet the individual tail feathers show the same antero-posterior
and hemilateral types of manifestation of secondaiy sex characters that
we find in the body as a whole.

The first type is illustrated by the hemilateral male and female
pigmentary pattern in each feather in my hermaphrodite specimen.
The second ty|5e is illustrated by the transverse distribution of the
same characters in the tail feathers in Shattock's specimen (cf. PI. XI,
fig. 14). Both represent extreme conditions of the normal arrangement
in the tails of many birds in which not only does the outer or less
covered side of the tail featheis exhibit a different pigmentary pattern
to the inner side, but the distal portion inaj' be also marked off from
the proximal portion by transverse areas of different pattern.

The value of this specimen seems to lie in the light it may po.ssiblv
ihrow on the process of Sex Differentiation. Tiiis individual bird shows

C. J. Bond 213

a tendency to divide into two halves of different sex, a male left halt'
and a female right half, and along with this somatic differentiation the
sex gland itself also shows a tendency to divide into a female portion
and a male portion. Owing further to the fortunate fact that these
two sex glands do not functionate coincidently, we have an opportunity
of observing a natural experiment, in which the effect of a female
Hormone is first exercised simultaneously on the male and the female
side of one and the same individual, while at a later stage the male
Hormone exerts its influence on the same dual organism, with the
result that while in the case of the trunk and limbs the male side only
responds to the male Hormone, the tail responds as a whole, but only
as regards one side of each individual tail feather.

Moreover along with, but not as the cause of or consequence of
this abnormal somatic segregation, anothei' abnormal segi-egation also
occurred in the sex gland by which a dual gland or ovo-testis was
formed and in which (as usual) the female portion came into ftinctional
activity and retrogressed before the male portion began to develop.

In sorne such manner it seems possible to explain the limitation of
influence of the male and female sex gland Hormone to opposite sides
of the body, for the side which possessed only femaleness and no niale-
ness (i.e. no somatic rudiments capable of developing male characters)
could not respond to the male Hormone, while the side which con
tained no female rudiments could not respond to the female Hormone.
There is however a difficulty in this explanation, for as I have already
pointed out (cf PI. XI, fig. 6), a well-marked spur rudiment such as
normally occurs in the hen phea,sant also occurs on the right leg (that
is on the female side) in this bird. How are we to interpret this fact ?
It is true that it is small and resembles the rudimentary spur in normal
female birds. Hence it may be regarded as a " specific " and not as a
" sexual " character and so may be considered to be out of the reach of
the influence so to .speak of the male Hormone.

Or it may be that this spur rudiment on the right, or female, side
represents a single dose of maleness instead of the double dose on the
left side, in which as a consequence the spur is well developed. If this
be so, then instead of simple " Presence and Absence " of malene.ss and
femaleness the question becomes one of the volume of these factors.
The greater volume of the male factor on the left side would then
account for the readier response to the stimulus of the male Hormone
in the left leg.

But whether we try to explain the facts by the presence and

214 Secondary Male Cliaracterfi in a Pheanant

absence of any given factor, or by the TuieLjiial volnine of any factor
on opposite sides of the body, or by the inhibition of one factor by
another, or by a third or metabolic fiictor as suggested by Geoffrey
Smith {Proc. Linnean Soc. March 6, 1913) and Doncaster (Brit.
Assoc. Birmingham, 1913), which acts as an intermediary between
the sex gland and the secondary sex characters, still this bird
differs from the normal individual in the fact that a process which
normally affects both sides of the body equally, in this case affects
them unequally. And yet this does not occur in a sharply defined or
clear cut manner. In some parts, as for instance the limbs, the head
and neck, this hemilateral limitation of influence is well marked, while
along the middle line of the body and in the postei'ior part of the trunk
there is some overlapping, and in the tail the influencing factor is
broken up into secondary factors which control individual tail feather
areas.

This latter is I think a point of some importance. It suggests that
the segregation of controlling factors (whatever may be their material
basis in heredity) does not occur suddenly at one stage only in the
development of the organism but that it runs on and accompanies
those cell divisions which at different stages regulate the growth of
the different organs and tissues of the body.

Now if the ontological process be continuous and gradual, may not
the segregation of gametic factors in the germ cell be a gradual and
continuous process also ? Although we draw an arbitrary line at the
stage of fertilization, it is probable that gametic and blastomeric
segregation are one and the same process which becomes accelerated
at the time of fertilization.

If this be so, then from what we know of the irregular and
asymmetrical cell divisions which lead to overlapping and blurring of
unit characters in the zygote, it is reasonable to suppose that those
qualitative divisions of the nuclear hereditary material (whatever it be,
by which the gametic factors, the precursors of unit characters, are
separated out in the germ cells) may also be less sharply defined than
we have been led to think. Segregation may not be the cutting
asunder of a chromosome or a portion of a chromosome by a sharp
division into two distinct portions but the pulling asunder of one
portion from another with detached particles adhering to either end.

C. J. Bond 215

Suggested explanatory hypothesis.

The question now arises whether any explanation can be suggested
which will fit the facts in this and similar abnormal cases. Assuming
that the fertilized egg fi'om which this zygote developed was a female
egg, i.e. heterozygous in respect of sex, then we must suppose that at
the stage of blastomeric segmentation at which the division of the body
into two halves was laid down, the se.x factor (instead of passing in
equal portions to each half of the germ) divided unevenly, the factor
for raaleness passing into the left half and the factor for femaleness
into the right half of the body. This lateral segregation was not
however clear cut in those rudiments which represent the median
fusion body areas, while in the tail region this qualitative segmentation
was delayed till those cell divisions occurred which control the develop-
ment of the separate halves of each tail feather.

The essential feature in this explanation is that it takes account
of genetic as well as physiological factors. It ascribes an abnormal
physiological phenomenon like the asymmetrical assumption of secondary
sex characters to the interaction of two factors, a peripheral or somatic
factor, and a central or sex gland gametic factor, though of course both
these owe their abnormal limitations to a common abnormality of genetic
origin, possibly arising at the time of the segregation between somato-
plasm and reserve germplasm in the zygote.

In thus falling back on genetic factors one is forcibly reminded of
certain features in modern views of heredity. There also the associa-
tion between the gametic factor in the fertilized ovum and the unit
character in the zygote is pictured in just the same way. The
gametic factor stands to the unit chai-acter in the same relation that
the sex gland and its Hormone stand to the corresponding secondary
sex character. Just as we are still unable to decide whether the
appearance of any given secondary sex character is due to the presence
of a corresponding sex gland Hormone, or to the absence of an opposite
sex gland Hormone which, if present, might inhibit the influence of the
former, so students of heredity are still unable to decide whether any
dominant unit character, such as tallness in the pea, or an abnormality
like Brachydactyly, depends on the presence of a factor for tallness or
short fingeredness, or on the absence of another factor which, if present,
might inhibit the action of this abnormal factor, and so make for
normality.

210 Secovdary Male Cliaractevfi hi a Pheamnt

In fact the " Presence and Absence " theory, the " Inhibition "
theory, and the "Unequal Vohmie" theory of Dominance and Recessivity
are all as relevant to the study of the physiology of sex as they are to
the study of the origin of all unit characters. Moreover, sex itself being
a unit character, observations on abnormal examples like the present
may possibly throw light on the larger pi-oblem.

For instance the fact that in vertebrates the sex gland influences
the epithelial or other tissues which develop secondary male characters
by means of some internal secretion, probably an enzyme, suggests that
genetic factors in the nucleus of the fertilized ovum exei'cise their long
drawn ciut influence over the development of all unit characters in
the zygote in the same way. In both cases three elements are con-
cerned :

((/) Peripheral cells or unit characters.

(h) Central sex gland or gametic factor.

{c) Some communicating medium, possibly an enzyme, circulating
in the blood rather than any structural continuity between (a) and (6).

We may even extend Ehrlich's side chain theory of immunity to the
field of the development of secondary sex characters. We may conceive
of the sex gland as liberating an Hormonic key capable o'l fitting the
peripheral cell lock, but this must be made available and carried to
the cell by some intermediate agency, and this intermediate handle
or amboceptor may be supplied by one or more of the other iluctless
g-lands.

Just as it is possible by cross fertilization to replace one gametic
factor by another and so to substitute one unit character for another
in the zygote, so also by the removal of the sex gland in the indi-
vidual (and more especially by parasitic castration) it is possible to
bring about the replacement of the secondary sex characters normal
to one sex by those peculiai- to the other.

JOURNAL OF GENETICS, VOL. IIL NO. 3

Fig. 1. Dorsal Surface. Hermaiiluodite
Pheasant.

Fig. 2. Ventral Surface. Heiuiapljrodite
Pheasant.

PLATE K

Fig. 3. Left or Male side. Hermaphrodite
Pheasant.

Fig. 4. Eight or Female side. Hermaphrodite
Pheasant.

JOURNAL OF GENETICS, VOL. 111. NO. 3

PLATE XI

I^^^H

^^^^B ' ?^^k^^l

W--

B^^^^^z^B

.^'"iv.^yf

^^^K^^^l

•Mi

^^^^^^Bpj^^^^^^^^l

■S--^-

^^

^

1

%

Fig. 5. Eight leg. Left leg with spur.
Hermaphrodite Pheasant.

Fig. 0. Kight leg. Enlarged. Kiidi-
laeut of spur. Hermaphrodite Pheasant.

Fig. 14. Tail feather of Cock Pheasant assuming Female colour patteru at proximal
or growing end. Transverse limitation of i and J characters. Specimen in
Museum Royal Coll. Surg. See Shattock, Trans. Path. Soc. Vol. lvii. (Copied
by permission.)

JOURNAL OF GENETICS, VOL. ML NO. 3

PLATE XII

Fig. 7. Unilateral male colour pattern in individual tail
feathers. Hermaphrodite Pheasant.

Fig. 8. Bilateral male colour pattern in individual
tail feathers. Hen Pheasant, atroph.y of ovary, and
presence of male elements.

Fig. 9. Cock Pheasant. Normal male
colour pattern in tail leathers.

JOURNAL OF GENETICS, VOL. IN. NO. 3

PLATE XIII

Fig. 10. Sex-glau.l. Ovo-testis (x2).
Hermaphrodite Phenstant.

Fig. 13. Sex-gland. Hen Pheasant. Symmetrical
assumption of male plumage, atrophy ovary.
A, Male portion. B, Female portion of gland.

Fig. 11. Ovarian portion of Sex-gland.
Hermaphrodite Pheasant.

Fig. 12. Testicular portion of Sex-gland.
Hermaijhrodite Pheasant.

MINOR-BRACHYDACTYLY. No. 2.
By H. DRINKWATER, M.D., F.R.S. (Edin.), F.L.S.

In September 1911, at a Meeting of the International Conference
of Genetics, held in Paris, I gave an account of a family showing
an inherited abnormality, which I termed Minor-Brachydactyly, to
distinguish it from a more marked defect of the same kind, termed
Brachydactyly , described previously^ The account of the Minor-
Brachydactyly family was also published in the Journal of Genetics,
Vol. II. Part I. February 1912. More i-ecently, owing to the courtesy
of my friend Dr F. Drinkwater of Llangollen, I have been able to study
another family showing the Minor-Brachydactylous condition. So
closely do the two families resemble one another, as regards this
abnormality, that one cannot help thinking they must have arisen
from a common stock, though the connection cannot now be traced,
and as far as the records go back there is no blood-relationship between
them.

The most marked peculiarities in each family are the shortness of
the digits (fingers and toes), and the shortness of stature.

As a full description was given in the accounts already referred to,
it will only be necessary in this paper, to draw attention to the main
features.

In the Brachydactyly family, the abnormality in the fingers was
shown to consist in a marked shortening of the middle phalanx, which
in the adult was found to be united to the terminal one : so that the
finger has two bones instead of the normal three. The length of the
finger was found to be about half the normal length.

In the Minor-Brachydactyly family there is also a shortening of the
digits, but to a smaller extent and due to the same abortive condition

1 Account of a Brachydactylous Family, Vroc. Buij. Sue. of Edin. Vul. xxviii. Fart 1.

218

Minor- Brachydactylff

-\i

o-

■CH-gM-

,o

<

o

- CH-

- CH

-^D— o

■\3-

II -

-0+

II

^3-

■^D

-O

CH--

\)

-^

-Of
-Of -

-\)

-CM-

♦4
■0+

-"o

-CH-
-CH-

-\)

-0+
-Of

\5

L\)

W

be

'e

o

II. Dkinkwatkk 219

of the middle bone, the second phahuix, which however always remains
in the adult separate from the terminal bone.

This second family of Minor-Brachydactyly can be traced through
five generations, and there are eight abnormal individuals alive at the
present time. These are ninubered in the chart, 4 to 11 inclusive.
I am able to present radiographs of each case, and photograjths of the
right hand of four of them.

Radiographs of the hand.

(1) Adults. These show the shortened middle phalanx in every
case. It remains a separate bone. (PI. XIV, fig. 1.)

(2) Children. The epiphysis is seen to be absent as a rule.
(PL XV, fig. 1.)

Radiographs of feet.

(1) Adults. Ankylosis has occurred between the second and
terminal phalanges in every case. The feet are therefore more de-
generate than the hands. (PI. XIV, fig. 2.)

(2) Children. The epiphysis is absent fl-om the base of the second
phalanx in every toe (except the big toe). (PL XV, fig. 2.)

Photographs of hands.

A comparison with the normal hand will show the peculiarities.
(PL XVI, figs. 1 and 2.)

Mendelism.

The abnormality behaves as a Mendelian dominant ; only being
reproduced by an affected individual. The children of normals are all
free from the abnormality and have fingers and toes of the ordinary
type. The expected ratio is approached as nearly as possible, for of
19 descendants of abnormals 9 are abnormals.

It is necessary to explain one part of the pedigree chart.

No. 1, a normal woman, and the female ancestor of all the abnormals,
was married to a normal man. All their descendants, as well as her
brothers and sisters, are seen to have been normal. In the village in
which this couple lived was a short-fingered man (No. 2) who was
occasionally employed by them to perform a certain surgical operation
on their young pigs. Eventually the woman gave birth to a daughter
(No. 3) who was Brachydactylous.

220 Minor- Braehiirlactylji

.imber

Age

4

—

5

3!)

6

32

7

28

9

7

10

91:

11

fi

Measurements in Inches.

Middle finger Hand width Height

2i 5J 37

2^ 6^ 60

2 53 59

•2\ 6| 59

2 i\ 43

If 4| 46f

If 44 424

The average length of the iniddle finger in the adults is 2\ inches:
about 1 inch shorter than the normal.

The average height of the adults is 58| inches : about 4^ inches
below the normal average.

EXPLANATION OF PLATES.

PLATE XIV.

Fig. 1. Right hand of woman showing minor-brachydactyly. No. 7 in pedigree.
Fig. 2. Right foot „ „ „ „

PLATE XV.
Fig. 1. Right hand of girl showing niinor-brachydactyly. No. 11 in pedigree.
Fig. 2. Right foot

PLATE XVI.
Figs. 1 and 2. Hands of mother and daughter. Nos. 4 and 6.

Fig. 3. Boy (No. 8 in pedigi'ee, aetat. 14) on left, and jounger normal brother, aelat. 12,
on right.

JOURNAL OF GENETICS, VOL, IIL NO. 3

¥,■■. 1.

PLATE XIV

!£JSS.2i;i^iaxiiii

Via. ■2.

JOURNAL OF GENETICS, VOL. IH. NO. 3

Fi-. 1.

FiH.

PLATE XVI

Fig. 3.

PEIMARY AND SECONDARY REDUPLICATION

SERIES.

By p. G. bailey, M.A.

(Clare College, Cambridge.)

In a recent paper, Trow (4) discusses theoretically the possible

interactions of the factors making up a three-factor group of such

a nature that any two may form a reduplicated series. He shows

that if the three factors of such a group have the primary reduplication

series

l:\:\:l

in : \ -.1 : m

■H : 1 : 1 : n,

then the secondary or observed reduplication series will be

Imn + I
linn + m
Imn + n

m + II : m + n : Imn + I
I + n : I + n : Im ii + m
I +m : I -\- m : Imn + n.

He further points out that if n becomes 1, that is if there is no
primary reduplication series between one of the pairs, the series become

m
Im + 1

1
1

/+ m

1 /

1 : m
I + m : Im + 1 .

For the convenience of reference the former series will be called Trow's
general hypothesis ; the latter series Trow's special hypothesis.

There exists, also, the possibility to which Punnett (3) calls
attention, namely that the primary reduplication series, obtained
by analysis as above of the facts observed when three factors are
involved, differ from the reduplication series found when only two
factors are involved. In order to avoid confusion it is desirable that
the three possible series should be given distinctive names. In this

Journ. of Gen. in 15

222 Primar]f ami Secondarji RedupUcation Series

paper the series found when only two factors are involved will be called
the ' fundamental ' series for those two factors ; following Tmw tjie
series observed when thi-ee factors are involved will be called the
secondary series : the underlying series calculated fi-oni these secondary
series will be called the primary series.

The data obtained from crosses involving three factors have been
discussed by other writers from the point of view of the special
hypothesis. Owing to the great importance of the subject of coupling
and repulsion it seemed to me that it would be interesting to study
the observed phenomena by means of the general hypothesis also.

The most complete account of such crosses is given b}' Punnett(3)
in a recent paper. The following calculations are based upon the
data there discussed, and reference should be made to that paper for
an explanation of the symbols used in the following account.

(a) Nature of mating EBL x ebl.

The equations from which the primary reduplication series can be
determined are

linn + / _ 13
VI + n 3 '

Imn + w- _ „„

I + n
Inui + n 13

I +m ~ ¥ '
which give

/ =)/= 2-1,

„, = 48-31.

These numbers indicate the complicated primary series

BL : Bl : bL : bl = 21 : 10 : 10 : 21

BE : Be : bE : be = 483 : 10 : 10 : 483

EL : El : eL : el = 21 : 10 : 10 : 21.

These complicated series however approach closely to the simple
series

2

: 1

: 1

: 2

r>2 :

: 1 :

: 1 :

52

2

: 1

: 1

• 9

' It may be noted that if the two smaller observed reduplication series are equal,
the assumption that n = 1 cannot be true, for then / = 1. However it is impossible at present
to decide whether these two series are in fact equal or not owing to the relative sraallness
of the numbers.

p. (Jr. Bailev 223

which would give rise to the following secondary gametic series:

39 : 10 : 10 : 89
65 : 1 : 1 : 65
39 : 10 : 10 : 39.

These gametic series would give rise to zygotic series, which agree
fairly closely with those actually observed, cf Punnett (3), p. 81 :

BL ; 81 : bL : bl
BE : Be : bE ; be
EL : El ; eL ; el

479 : 58 : 66 : 143 observed

4S0 : 68 : 68 : 1^2 cede.

532 : 5 : 6 : 203
■5o4- : '5'/' : o'7 : 184

479 : 59 : 66 : 142

A9U : 68 : 68 : J J.J.

(/9) Nature of mating BeL x bEI.

The observed BL relationship is most accurately explained on the
basis of a 10 : 1 : 1 : 10 series, and the EL relationship on a 1 : 12 : 12 : 1
series, but it is by no means impossible that these gametic series are
m reality of the same intensity. It will be assumed for the sake of
simplicity that they are.

The equations may then be written

111 + I
l-in + ni

>32.

11

The only value of / which would be of a simple nature and would

approximately satisfy these equations is Z = 3. Then the observed

secondary relations between B and L and E and L must be of a type

with less intensity than 9 : 1 and greater intensity than 8:1.

The observed (cf (3), p. 83) and calculated zygotic series are given

below :

3006 : 164 : 212 : 843 observed
BL:BI:bL:bl;: ^^^^ , ^^^ , ,^^ . ^.^ ^^^^_ 9 : 1 : 1 : 9 basis

2200:1001:1018: 6 observed
el :: ^^^. , ^^^^ . ^^^^ . ^y.^. ^^^^^.

'2'24: Priinarii and Secondar;/ Reduplication Series

(7) Nature of mutinrj DfN x dFn.

The value of L is again very near that of n if not equal to it.
Assuming I = n, the observed secondary reduplication series can be
shown to depend upon the complicated series

19 : 10 : 10 : 19
10 : 57 : 57 : 10
10 : 19 : 19 : 10.

These complicated series are not very different from the simple series

2:1:1:2
1:6:6:1
1:2:2:1,

which would be obtained if the observed secondary series were

3-2:1:1: 32
1 : 7-3 : 7-3 : 1
1 : 3-2 : 3-2 : 1.

Below are given for purposes of comparison the actual numbers
obtained (cf. (3), p. 89), the numbers to be expected upon the above
hypothesis, and the numbers to be expected on Trow's special hypo-
thesis.

Expectation on Trow's si^cial hypothesis

^

On the assumption
secondary

tliat this is tlie
series

Found

Expectation on

3-2:1:1: 32

system

On tlie assumption

that this is tlie primary

series 3:1:1:3

If tlie DF series
is 7:1

If the DF series
is 15 : 1

ND

282

386

284

273

277

Nd

49

46

48

58-3

54

nD

52

46

48

58-3

54

nd

59

65

On

1:3-2: 3-2 :1

62
On

1:3:3:1

52

56-5

NF

22.5

^^}

228

231

230

Nf

106

lOi-2

103 -.5

100

101

nF

101

104-2

103-5

100

101

nf

10

6-3

Ou

1 : 7-3: 7-3:1

7

On

1:7:7:1 1

On

. : 15 : 15 :

10-8

1

9

DF

220

323

222-7

221-4

Df

114

109

108-8

110-1

dF

106

109

108-8

1101

df

2

1-6

1-7

-4

p. G. Bailby 225

The above table shows that Trow's special hypothesis fits the
figures better than the general hypothesis', if the assumption be made
that the NF series is the secondary' series, and that the DF relation-
ship is on the 1 : 15 basis. The agreement may be purely accidental
but it is interesting to note that Pimnett found certain strains in which
the fundamental N F series was itself on a 1 ; 1 basis.

(fi) Nature of mating DFnxdfN.

The values obtained in this case are l = n = l, and m= 15, i.e. the
apparent relations are the real relations. The N D and N F fundamental
repulsion series are reduced to a 1 : 1 : 1 : 1 basis.

The only additional data- bearing upon this question are those
furnished by Gregory (2) in his description of the results obtained
from a cross of the nature MSG msg x msg msg. Trow (4), p. 315,
discusses these results from the point of view of the special hypo-
thesis, and he shows that the observed numbers agree fairly well with
those obtained by calculation on the assumption that the SG series is
the secondary one. On the other hand it can be shown that, if in
reality the general hypothesis applies to this case, the numbers fit in
very badly with any simple primary series. It should be noted that
the observed MS series closely approximates to the fundamental MS
series, and that no fundamental SG series has yet been described.
Consequently it is by no means clear that the case, which Gregory
has described, is really comparable with those described by Punnett.

Co7iclusion.

The general hypothesis adopted above, although it admits the
possibility of a difference between the fundamental and the primary
series due to the interaction of the reduplication series one upon
another, does not postulate a differential interaction. The special
hypothesis on the other hand does postulate a differential interaction.

1 If the complex series calculated on the general hypothesis be taken, the agreement
between the numbers found and the numbers obtained by calculation would be even better
than on Trow's simple hypothesis.

2 Morgan and Cattell (5, 6) have described certain crosses with Drosophila which
involve three factors. The results, however, are complicated by the phenomena of sex
limitation, and by differential death rates. Moreover it is not clear in each case whether
the given relationships are to be looked upon as fundamental or primary. Nevertheless
it is interesting to note that in the most satisfactory case, namely that involving black
body colour B, red eye colour R, and long wings L, the secondary relationship for Land B
i.e. 1-9 : I calculated upon Trow's special hypothesis closely approximates to the relation-
ship found by experiment.

'226 Priinar// and Secondunj Reduplication Series

For three factors taking part in a reduplicatidii series, Bateson and
Punnett (1) suggest an octant arrangement. 8uch an octant arrange-
ment showing the possible course of the divisions in the formation of
reduplication series is given below.

imn

Imii^^,^

r~--l

X/^

?x

/ X"^

T / \

M*o\j

{^\

U^y^

V.^-c /

\ / o

''-X /'

\/ «

'^\/

X. '5'

It is difficult to see any cause for differential treatment on such
a scheme. Trow's scheme, however, which involves one factor waiting
its turn to segregate until the other two have completed their redupli-
cation series, does from its very nature offer a jjossible explanation for
such differential treatment. Consequently further research showing
whether the course of events is best explained by the general hypothesis
or by the special hypothesis may throw light upcjn the question as to
whether the octant scheme or Trow's scheme is to be preferred as a
better picture of the process of segi'egation and the formation of
reduplication series.

An interesting feature that becomes apparent on analysing the
observed fact by means of the general hypothesis, is the regularity of
the underlying phenomena. Such analysis shows that in all four
cases the two fundamental series of least intensity have their intensity
reduced when they become primary series, although the observed or
secondary series may be of a greater intensity. It is by no means
impossible that the same holds good for the third reduplication series,
as is certainly the case in the EBL x ebl and in the DfN x dFn
uiatings.

Another possible regularity in the observed phenomena is the
reduction of two of the fundamental series to primary series of
identical intensity. The numbers are strongly in favour of this sug-
gestion in at least two of the cases. If such a relatitmship is shown
to be general, it may be due to the necessity for the divisions involved
in segregation and in the formation of reduplication series to be on
a symmetrical jilan.

p. G. Bailey 227

LITERATURE.

1. Bateson, W., aud Punxett, R. C. " On Gametic Series involving Redupli-

cation of certain Terms." Journal of Genatics^ 1911.

2. Gregory, R. P. "On Gametic Coupling and Repulsion in Primula sinensis."

Proc. Roy. Soc. 1911.

3. PuNXETT, R. C. "Reduplication Series in Sweet Peas." Jonrnal of Genetics,

1913.

4. Trow, A. H. " Forms of Reduplication, Primary and Secondary." Journal

of Genetics, 1912.

5. Morgan, T. H. and E. Cattell, 1912. " Data for the Study of Sex-linked

Inheritance in Drosopkila." Journ. E.vp. Zool. xili.

6. , 1913. " Additional Data for the Study of Sex-linked Inheritance in

Drosophila." Journ. E.rp. Zool. xiv

THE

CAMBRIDGE BULLETIN

No. XXX

October MCMXIII

Cambridge : at the University Press

The

Cambridge Bulletin will in

future be

arranged in three parts:

I.

An illuitrated descriptive account

of recent books.

II.

A classified list of all books pub-

lished since the last issue of the

Bill If tin.

III.

Announcements of forthcoming

books.

Copies of the Bulletin will be regularly

sent, post

free, to any address on applica-

tion to M

r C. F. Clay, Cambridge Univer-

sity Press,

Fetter Lane, E.C.

d^ THE CAMBRIDGE BULLETIN ^*?

No. XXX

October 191 3

Collected Literary) Essayis. Classical and Modern. By A. JV. Verrall^
Litt.D., King Edward Fll Professor of English Literature and Fellow
of Trinity College, Cambridge, Hon. Litt.D. Dublin. Edited by M. A.
Bayfield, M.A., and J. D. Duff, M.A.

Demy 8vo. pp. cxiv + 292. With a memoir and a portrait. Price lOJ. (>d. net

Collected Studies in Greek and Latin Scholarship. By the same author and
editors.

Demy 8vo. pp. viii + 372. Price loi. 61/. net

Not the least interesting portion of the first of the abo-\'e volumes
is the Memoir, in which the following
passages illustrate two of Verrall's char-
acteristics— his Liberalism and his love
of nonsense: "And such a Liberal was
Verrall, as he himself used to say.
Miss Jane Harrison tells a confirmatory
story : —

I remember saying to him apropos of some
scholar from whom I differed, 'It is intolerable
that people should be allowed to go on talking
and teaching such nonsense!' He screwed up
a whimsical eye at me and said, ' All right,
let's have back the Inquisition.' "

" But the joy of joys was his manner
of reciting humorous verse or pure non-
sense, and to find (if it was your first
experience of him in this vein) that he
took as intimate a delight in it as you
did yourself. 'Tragedy!' he once
said to me suddenly in the early days ;
' Did you ever hear this .' ' And he
proceeded to chant slowly, in rolling, melancholy tones, a once famous
song of Toole's (metre strictly dactylic) —

'A norrible tale I 'ave to tell

Of the sad di-sasters that befell
A noble family as once re-sided

In the very same thoroughfare as I did.' "

The Literary Essays include amongst others : The Feast of Saturn,
A Villa at Tivoli, The Birth of Virgil, Aristophanes on Tennyson, The Prose
of Walter Scott, and '■'■Diana of the Crossways." Of the two last-named
The Times says that " they are not only the best essays in the collection,
which is saying much, but they are also, beyond comparison, the best
essays on their subjects."

I I 2

The late Dr Verrall

GREEK SCYTHIA

Scy)th(dns a.nd Greeks. -^ survey of ancitnt bhtor\ and archaeology on the
north coast of the Eiixine from the Danube to the Caucasus. By Ellis H.
Minns., M.A., Fellow of Pembroke College., Cambridge., Member of the
Imperial Russian Archaeological Society.

Royal 4to. Buckram, gilt top. pp. xl + 720. With 9 maps and plans,
9 coin plates, and 355 illustrations in the text. Price £2. 3J. net

This book offers a summary of what is known as to the archaeology,
ethnology and history ot the region between the Carpathians and the
Caucasus. The region is of varied importance for different branches
of knowledge touching the ancient world, yet about it the scholars of
Western Europe have had a certain difficulty in obtaining recent information,
because each found it unprofitable
to master Russian for the sake of
pursuing his subject into an out-
lying corner. The language diffi-
culty, therefore, first suggested this
work, and the author's original
intention was merely to supply a
key to what has been written by
Russian scholars. But such a
fragmentary account of things
would have been most unsatis-
factory, and enough advance has
been made since the last attempt
to review the subject, to justify a
provisional summary.

Though thegeographical limits
have confessedly been dictated by
considerations of language, yet
the frontier of Russia towards
the Carpathians and the Danube
answers nearly to a real historico-
geographical boundary, the western
limit of the true steppe. The
Caucasus, again, is a world in itself, having little in common with the steppe,
nor has the time yet come to bring any sort of system into its archaeology.
On the other hand, the unity of the Asiatic and European steppe has led
the author on occasion right across to Siberia, Turkestan and China.

"The book," to quote the words of The Athenaeum, "is in itself
a library on Greek Scythia and we trust it will receive full recognition
both at home and abroad. The author's knowledge of Russian and his
intimacy with the sites he describes bring him constantly nearer to his
sources than most writers can hope to penetrate. For all these reasons
we commend his work both to the learned world and to educated men
of the world."

Fig. 58. Chmyreva Mogila. Golden
harness adornment. \.

ROMAN BOROUGHS— LATIN PRONUNCIATION

The Municipalities of the Roman Empire. By James S. Reid, Litt.D.,
Fellow of Gonville and Caius College, Cambridge, Professor of Ancient
History in the University of Cambridge, Hon. Litt.D. {^Dublin), Hon,
LL.D. [St Andrews).

Demy 8vo. pp. xvi + 548. Price 12J. net

The volume is the outcome of a course of lectures given in the
university of London and afterw^ards in America, and surveys the Roman
empire in its character of a vast federation of commonwealths, emphasising
the historical significance of the great movement of civilisation whereby
for loose rural and tribal unions was substituted a civic system, and the
importance in the annals of the Roman empire of the growth and decline
of the towns. The book is planned as a survey of the empire, province
by province, so as to shew how the Roman rulers influenced the develop-
ment and decay of the municipal system in each.

"Our survey," says the author in Chapter xv, "has shewn us abundantly
that something of the dignity of sovereignty hung round the ancient city
down to a late age, and that this colours ancient municipal institutions and

differentiates them profoundly from their present-day counterparts The

inhabitants of the territory of each municipality were in a way a little
nation, whose affections were mainly centred in the town where its public
affairs were carried on, its festivals celebrated, and its gods revered. Every
man aspired to have a domicile within the walls if he could, and all those
who performed public functions were compelled to reside there or within

a thousand paces, as a rule It was this association of the burgesses

en masse that constituted for the ordinary man the chief element in
well-being. The life within the home counted for infinitely less, the
life without the home for infinitely more than in modern times."

"An important contribution to the history of the Roman Empire from a point of
view novel to the ordinary student ; at the same time it makes alluring reading, by
reason both of the freshness and lucid arrangement of the subject-matter, and of the
picturesque style in which the story is told." — Athenaeum

Quantity) and Accent in the Pronunciation of Latin. By F. IF. IVestaway.

Crown 8vo. pp. xvi + iii. Price 3/. net

In the Preface the author makes a spirited attack upon " the remnant
of the old school " which still clings to its "duU'sy dome'um [diilce damiim)
and its " nice-eye pry-us " {nisi prius) and addresses his book not to school-
masters but (i) to private students who desire to learn to pronounce and to
read Latin correctly, and (2) to those who feel that their acquired pronun-
ciation needs overhauling.

3

FOUNDATION OF THE WESTERN EMPIRE

The Cambridge Medieval History. Planned hy J. B. Bury, M.J., Regius
Professor of Modern History. Edited i>y H. M. Gwatkin, M.A., and
J. P. JVhitney, B.D. Volume II, The Rise of the Saracens and the
Foundation of the Western Empire.

Royal 8vo. pp. xxiv+Sgo. With a portfolio of maps.
Price 20J. net; to subscribers 15s. net

This volume covers the stormy period of about three hundred years
from Justinian to Charles the Great inclusive. It is a time little known
to the general reader, and even students of history in this country seldom
turn their attention to any part of it but the conversion of the English.
Hence, English books are scarce — Dr Hodgkin's Italy and her Invaders is
the brilliant exception which proves the rule — and the editors have had
to rely more on foreign scholars than iji the former volume. Some, indeed,
of the chapters treat of subjects on which very little has ever been written
in English, such as the Visigoths in Spain, the organisation of Imperial
Italy and Africa, the Saracen invasions of Sicily and Italy and the early
history and expansion of the Slavs.

The contents are as follows :

'Justinian. The Imperial Restoration in the TVest (Professor C. Diehl).
Justinian s Government in the East (Professor C. Diehl). Roman Laiv (Dr
H. J. Roby). Gaul under the Merovingian Franks. Narrative of Events
(Professor C. Pfister). Gaul under the Merovingian Franks. Institutions
(Professor C. Pfister). Spain under the Fisigoths (Dr Rafael Altamira).
Italy under the Loynhards (Dr L. M. Hartmann). Imperial Italy and
Africa : Administration (Dr L. M. Hartmann). Gregory the Great (the
Ven. Archd. W. H. Hutton). The Successors of Justinian (N. H. Baynes).
Mahomet and Islam (Professor A. A. Bevan). The Expansion of the Saracens.
The East (Professor C. H. Becker). The Expansion of the Saracens. Africa
and Europe (Professor C. H. Becker). The Successors of HeracHus to 717
(E. W. Brooks). The Expansion of the Slavs (T. Peisker). Keltic
Heathenism in Gaul (Professor Camille Jullian). Keltic Heathenistn in
the British Isles (Professor Sir Edward Anwyl). Germanic Heathenism
(Miss B. Phillpotts). Conversion of the Kelts. Roman Britain. Ireland.
Scotland (the Rev. F. E. Warren). Conversion of the Teutons. The
English. Germany (the Rev. Professor J. P. Whitney). England (to
c. 800J and English Institutions (W. J. Corbett, M.A.). The Carlovingian
Revolution, and Frankish Intervention in Italy (Professor G. L. Burr).
Conquests and Imperial Coronation of Charles the Great (Dr G. Seeliger).
Foundations of Society (Origins of Feudalism) (Dr P. VinogradoflF). Legislation
and Administration of Charles the Great (Dr G. Seeliger). The Papacy, to
Charles the Great (the Rev. F. J. Foakes-Jackson).

THE EASTERN QUESTION— ANCIENT WESSEX

The Ottoman Empire, t80t—t9t3. By William Miller, M.A. [Oxon),
Hon. LL.D. in the National University of Greece, Corresponding Member
of the Historical and Ethnological Society of Greece, author of The Latins
in the Levant.

Crown 8vo. pp. xvi + 548. With 4 maps. Price yj. 61/. net

The latest volume in The Cambridge Historical Series "could not,"
says The Westminster Gazette, "have been published at a more opportune
moment." It has been based wherever possible upon original docu-
ments, and traces the history of the Near East from the beginning of the
Nineteenth Century to the outbreak of the Balkan War and the Balkan
Conference at London in March 1913 ; its scope may be seen from the
following; list of contents :

Tile Ottoman Empire at the Dawn of the Nineteenth Century.
Napoleon in the Near East (1801-15). The Servian Risings (1804-1 7).
The Preface of Greek Independence (1815-21). The War of Greek
Independence (1821-9). The Creation of the Greek Kingdom (1829-33).
The Balkan and Syrian Difficulties of Turkey (1822-45). Greece under
the Bavarian Autocracy (1833-43). The Greek and Ionian Constitutions
(1843-53). The Crimean War (1853-6). The Union of the Danubian
Principalities (1856-62). The Cession of the Ionian Islands (1862-4).
Reforms and their Results: The Lebanon and Crete (1856-69). The
Roumanian and Servian Questions (1862-75). The Bulgarian Exarchate
(1870-5). The Balkan Crisis of 1875-8. The Union of the Two
Bulgarias (1878-87). Armenia, Crete, and Macedonia (1887-1908).
The Turkish Revolution (1908-12). The Balkan League (October
1912-March 19 1 3). Table of Rulers. Bibliography. Index.

Lord Cromer, writing in The Spectator, declares that " Mr Miller has
performed a most useful service in affording a guide by the aid of which
the historical student can find his way through the labyrinthine maze of
Balkan politics."

Earty} Wars of Wessex, Being Studies from EnglancVs School of Arms in
the West. By Albany F. Major, Author of Sagas and Songs of the
Norsemen, etc. Edited by the late Chas.JV. Whistler, M.R.C.S.,
Author of A Thane of Wessex, King Alfred's Viking, etc.

Demy 8vo. pp. xvi + 238. With 21 maps and plans. Price loi. 6</. net

A series of studies in the history of the pre-Norman period from the
foundation of the kingdom of Wessex in 495 to the campaign of 876-8.
Their distinguishing feature is that, while dealing only with the warfare
that took place in a comparatively limited area between Saxon and Briton
and Saxon and Dane, they show, nevertheless, that these " battles of the
kites and crows," as Milton contemptuously called them, had a far-reaching
influence on the future of England.

s

ANCIENT AND MODERN CHRISTIANITY— LATIN HYMNS

Paganism and Christianity) in Egyipi. By Philip David Scott-Moncrieff,
M.A. Cantab.

Crown 8vo. pp. x + 226. With a frontispiece. Price 6s. net

This little work was produced by a scholar whose sudden death in
February 191 1, before he had completed his twenty-ninth year, came as
a painful surprise to his many friends. He approached his subject from
the Egyptian point of view, he had a first-hand knowledge of the ancient
Egyptian religion, and as an archaeologist he was able to form his own
estimate of the bearing of the archaeological upon the purely literary
evidence, and in several cases to use his data in a novel way.

After a sketch of the condition of the old Egyptian religion at the
close of the Ptolemaic era, the beginnings of Christianity in Egypt are
discussed, and by fresh evidence from the papyri and the early Christian
tombs fresh light is thrown upon an obscure chapter of Church history.

"In the 200 pages of this small book," says T/'f iSia/xwrtw, "there will be
found more fresh matter than in many a bulky volume ; it will be received
as a valuable contribution to the study of the beginnings of Christianity in
the land of the Pharaohs."

The Interregnum. Twelve essays on religious doul>t. By R. A. P. Hill,
B.J., M.D.

Crown Svo. pp. xv+149. Price 31. 6J. net

The title of the book indicates "that stage in a man's mental
development when the old beliefs and sanctions of childhood are lost
and he has not yet had time to form new views of his own," and the
essays, written from the point of view of a man in such a condition,
endeavour to show first that it is a natural, right and reasonable thing for
a man who is striving to know the good, to sympathise with Christians
as a Christian during the interregnum, even though the balance of
evidence may seem to liim to be against Christianity ; and secondly that
rival systems have their difficulties and objections no less than Christianity,
and that the strength and weakness of materialistic theories can only be
realised by minds that are truly open and thought that is truly free.
Further, the author shows that there is reasonable ground for thinking that
Christianity is true from the purely speculative standpoint.

Earl}) Latin Hyimnaries. An Index of Hymns in Hymnaries before 1 1 00.
// ith an appendix from later sources. By 'James Mearns^ M.A.

Demy Svo. pp. xx-|-io8. With frontispiece. Price 5J. net

This index provides a convenient book of reference for those interested
in early Latin hymns and gives a fair idea of those actually used in Europe
before 1 100.

6

THE LITURGY— TWIN-CULTS— THE SONG OF SONGS

The Earlsi Histor}^ of the Liturgy. By J. H. Srawley^ D.D., Rector of
JVeeting, Norfolk, sometime Tutor of Selwyn College, Cambridge.

Small crown 8vo. pp. xx + 252. Price 6s. net

In this volume oF The Cambridge Liturgical Handbooks the word
"Liturgy" is used to denote "the order of service employed in the central
rite of the Christian Church, the Eucharist"; and Dr Srawley, after tracing
its history in the Apostolic and sub-Apostolic Ages, in Egypt and Alex-
andria, in Palestine and Syria, in the churches of Asia and Northern Africa,
in North Italy and Rome, treats in his last two chapters of the development
of the Liturgy and the early conceptions of the Eucharist which its history
illustrates.

Boanerges. By Rendel Harris.

Demy 8vo. pp. xxiv + 424. Price 15^. net

In this volume, in which the researches embodied in The Dioscuri in
Christian Legends (1903) and The Cult of the Heavenly Twins (1906) are
continued, the author shows the antiquity and wide diffusion of Twin-cults
and traces their modification from the primitive form which demands the
murder of the mother and children to the milder forms which lead to the
establishment of twin sanctuary towns.

It is made clear that these twin-cults have made a deep mark upon
ancient and modern religions, and that they affect the legendary sides of
the Old and New Testaments as well as the mythologies of Greece
and Rome.

The Song of Songs. Edited as a dramatic poem, luith introduction, revised
translation, and excursuses by If^illiam JValter Cannon.

Demy 8vo. pp. viii+i5S. Price yj. dd. net

The author's aim has been to write a book for those general readers
who would like to have some knowledge of the subject and hardly know
where to look for it. He wishes his readers not only to understand the
Song but to enjoy it as " the best Song of all."

" It is not often that one meets with so completely delightful and satis-
factory a work as Mr Cannon's study of 'The Song of Songs. '...Mr Cannon
gives us a full and impartial explanation of the traditional theory, according
to which the poem describes the progress of the mutual affection, and the
subsequent marriage, of Solomon and the Shulamite ; and also of the Syrian
Wedding theory, which has recently won the support of very eminent
scholars. According to his own view, the Song is a dramatic poem, not

intended, however, to be acted on the stage Mr Cannon's book is a most

beautiful study of a poem of consummate charm." — JVestminster Ga%ette

7 1—5

THE LERINS— SION COLLEGE

The History of the Islands of the Levins. The MonaUery, Saints, and
Theologians of S. Honorat. By A. C. Coopef-Marsdin^ D.D., Houorar\
Canon of Rochester.
Demy Svo. pp. vii + 336. With 15 illustrations. Price \os. 6J. net

This work follows on the author's Life of Caesarius, Bishop of Aries,
and, as an account of the leaders of Christian thought in the fifth and
sixth centuries who were trained in the monastery of S. Honorat, fills a

The Cell of "The Man with the Iron Mask"

gap in English theological literature. After reviewing the islands themselves,
the lives of S. Vincent and S. Patrick, and the foundation of the monastery,
the author treats of the monastery as a " Nursery ot Bishops," and in an
appendix deals witii Ancient Buildings, Treasures and Monuments.

Sion College and Library. By E. H. Peam\ M.J., Canon of ireitmlnster,
a Trustee of Sion Hospital, and formerly President of Sion College.
Demy Svo. pp. viii+374. With 4 illustrations. Price 91. net
"Perhaps those who will take most pleasure in Canon Pearce's book will be

such as have themselves walked about Sion But this well-written volume is one as

well for the general reader with historical or antiquarian tastes. Even dipped into here
and there in a spare hour, it will be found to yield much treasure to inform the mind,
to stir the imaijination with curious, incisive pictures of bygone days and manners, and

sometimes to evoke a hearty laugh Yet let it not be supposed for a moment that this

is other than a sound and exhaustive history of its subject." — GiiarJiaii

ENGLISH LITERATURE

The Cambridge History of English Literature. Edited by Sir A. W. Ward,
Litt.D., F.B.A., Master of Peterhouse, and A. R. Waller, M.A.,
Peterhouse. Volume X, The Age of "Johnion.

Royal 8vo, pp. xvi + 562. Price in buckram, 9;. net, in half-morocco,
1 5i. net ; to subscribers 7/. (>d. net and 1 2X. 6d. net respectively

As was hinted by the editors in the Prefatory Note to volume IX,
the canvas of English literature grows more and more crowded as the
eighteenth century approaches. The present volume is entitled The Age
of yohnson, and the contents are as follows :

Richardson (L. Cazamian). Fielding and Smollett (Harold Child).
Sterne, and the Novel of his Times (Professor C. E. Vaughan). The Drama
and the Stage (Professor G. Nettleton). Thomson and Natural Description
in Poetry (A. Hamilton Thompson). Gray (The late Rev. Duncan C.
Tovey). Toung, Collins and Lesser Poets of the Age of Johnson (Professor
George Saintsbury). Johnson and Boswell (David Nichol Smith). Oliver
Goldsmith (Henry Austin Dobson). The Literary Influence of the Middle
Ages (Professor W. P. Ker). Letter-Writers, ' I (H. B. Wheatley) ;
//, The Warwickshire Coterie (The Ven. Archdeacon W. H. Hutton).
Historians, I, Hume and Modern Historians (The Rev. William Hunt,
D.Litt.) ; //, Gil?i>on (Sir A. W. Ward). Philosophers (Professor W. R.
Sorley). Divines (The Ven. Archdeacon W. H. Hutton). The Literature
of Dissent, 1 660- 1 760 (William Arthur Shaw). Political Literature,
1 755-1 775 (C. W. Previt^-Orton).

"The essay on Gray is a masterpiece of analysis and criticism. ...Mr David Nichol
Smith admirably condenses many opinions of the ' Great Cham ' when he says that he
had 'a supreme talent for definition,' A better choice of critic for Oliver Goldsmith
than Dr Austin Dobson could not have been found ; while the essay on Gibbon by
Sir A. W. Ward is notable alike as a study of character and as an assessment of the
intrinsic value and wide influence of the 'Decline and Fall. '...The nation has good
reason to be proud of the plan and progress of The Cambridge History of English
Literature." — Manchester Courier

Outlines of Victorian Literature. By Hugh Walker, LL.D., and Mrs Hugh
Walker.

Large crown 8vo. pp. viii-l-224. Price 31. net

Based upon Dr Walker's larger volume. The same general plan has
been followed, but the scale reduced and care taken to write as simply as
possible ; above all, an attempt has been made to present the authors of
the period not merely as writers but as men. The six chapters of the
book are entitled Carlyle and the Systematic Thinkers, Poetry, Novels and
Novelists, The Historians, Biography and Criticism, The Fragments that
Remain. The Outlook recommends it to " all who are about to take up
for the first time the serious study of the literature of a truly notable
epoch."

THE RED SEA— THE GALEA TONGUE

Desert and Water Gardens of the Red Sea. Bang an account of the natives
and the shore formations of the coast. By Cyril Crossland, M.A. Cantab..,
B.Sc. Lond., F.L.S., F.Z.S.., Marine Biologist to the Sudan Government.

Demy 8vo.

pp.

xvi+158. With 91 maps and illustrations and
12 diagrams. Price 10s. 6J. net

The portion of the Red Sea coast described in this book (that between
18° N. and 22^ N. on the western side) is, says the author, one of the least
known coast-lines in the world. The country is a desert with a sparse

population ot nomads; no steamer passes
within miles ot the outermost reefs; and
native vessels sail by at the rate of about
one a month.

The fact, moreover, that the country
is only artificially made habitable at all adds
interest to the author's narrative in which
information of general interest, as well as a
description of things and peoples peculiar
to the country, is included.

Part I deals with the coast and its
people, social and religious conditions,
fishermen and pearl divers ; Part II with
corals and coral animals, the building of
reefs, and the making of the Red Sea.

In a postscript to his preface the
author finds that his book has a moral,
a thing never intended ! " It is that real romance and beauty are to be
found in things as they are, so that the man of science, popularly supposed
to be hardened by ' materialistic ' pursuits, has opportunities for a truer
worship than has the sentimentalist who bows before idols of his own
imagination."

Arabian Sword Dance

A Galla-English English-Galla Dictionary). Collected and Compiled by
E. C. Foot., F.R.G.S. Published with the aid and approval of H.M.
Foreign Office.

Demy

PP-

viii-H 118. Price 6.'

"The Galla," says the compiler in his Preface, "are a very numerous
people. They are found in Abyssinia from Harrar on the east to the Sudan
frontier on the west and from Wollo down to the southern frontier."

Sir John Harrington, for twelve years H.M. Minister in Abyssinia,
declares that "a book of this nature has been a long-felt want in the in-
terest of travellers and others to facilitate direct dealings with the Gallas.

It should be of service not only in Abyssinia but also on her frontiers

with the Sudan, Uganda and East Africa."

10

THE CHINA-TIBET FRONTIER

Ward, to whose memory

The Land of the Blue Poppy:. Travels of a luituraliit in Eastern Tibet.
By F. Kingdon IVard, B.A., F.R.G.S.

Royal 8vo. pp. xii + 284. With 40 plates from photographs by the
author and 5 maps. Price 12s. net

A son of the late Professor H. Marsha
the book is dedicated, the author
explains the raison d^etre of the
volume in his first paragraph :

" On my return from Western
China in September 1 9 10, I settled
down to humdrum life with every
prospect of becoming a quiet and
respectable citizen of Shanghai.
But.., travel had bitten too deeply
into my soul and.. .when after
months of civilized life something
better turned up, I accepted with
alacrity. This was none other
than the chance of plant-collecting
on the Tibetan border of Yun-nan."

The results of this expedition
are recorded in the book and a
preliminary list of 200 plants col-
lected (several of them being new
species) is given in an appendix ;
but Mr Ward studied men and
manners as well as plants. Here
is part of his description of a
Tibetan festival :

" The Tibetans always strike
me as being so much more jolly
and irresponsible than the Chinese.
...The children picked bunches of
flowers just as English children
love to do, romped, made swings,
and swung each other and finally

sat down to eat cakes In the

evening they all trooped back to

the village to dance in the mule

square and skip. Three or four

little girls would link ;arms and

facing another similar line of girls

advance and retreat by turns, two steps and a kick, singing a not

unmusical chorus — a most delightful parody of ' Here we go gathering

nuts and may '."

II

A Tibetan girl of A-tun-tsi

MAPS AND SURVEY— MADRAS

Maps and Survey. By A. R. Hinks, M.A., F.R.S., Ass'ntant Secretary to the
Royal Geographical Society, Gresham Professor of Astronomy.

Demy Svo. pp. xvi + 206. With 24 plates. Price 6s. net

This introduction to the study of maps and the processes of survey by
which they are made has been written to supply the need of a book giving
a general account of the many-sided art of survey, which his experience as
a teacher in the department of geography in the university of Cambridge
has shown to exist. The treatment of the topographical and geodetic
survey follows closely the methods employed by the Ordnance Survey,
the Survey of India, and the School of Military Engineering at Chatham.
The general scope of the book may be seen from the following list
of chapters : Maps — Map Analysis — Route Traversing — Simple Land
Survey — Compass and Plane Table Sketching — Topographical Survey —
Geodetic Survey — Survey Instruments.

" Witliout being a technical treatise on surveying, this book deals with the subject
in sufficient detail to put any inap-maker on the track of the methods best suited to his
purpose. ..though short and compact it is a book evidently based on a thorough know-
ledge of the subject and should interest a wide circle of students of geography." —

Scotsman

The Madras Presidency, with Mysore, Coorg, and the associated states.
By Edgar Thurston, CLE., sometime Superintendent of the Madras
Government Museum.

Large crown Svo. pp. xii+29+. With 100 maps and illustrations.

The first volume of the Provincial Geographies of India (see p. 32) deals
with the Presidency of Madras in its physical, ethnological, archaeological,
historical and industrial aspects.

In his Preface the General Editor, Sir T. H. Holland, says : "Among
the ' provinces ' the Madras Presidency has above all developed an in-
dividuality of its own — advanced in education through early missionary
effort, free of frontier worries, comparatively homogeneous in ethnic
composition, and sufficiently unknown to the Central Government to
escape undue interference, its officials and its people are distinctly
' Madrassi,' and are rightly proud to be so. No geographical unit could
more appropriately be selected to initiate this series, and everyone who
knows the Senior Presidency will recognise the pre-eminent fitness of
Mr Edgar Thurston to give a true picture of South India. As Superin-
tendent of the Madras Museum, he sampled every form of natural product
in the South. As Superintendent of the Ethnographic Survey he obtained
an intimate acquaintance with the people."

12

RUBBER AND RUBBER PLANTING

Rubber and Rubber Planting. By
Board oj Agriculture and Fisheries
Gardens, Ce\lon.

R. H. Loci, Sc.D., Inspector H.M.
sometime Assistant Director of Botanic

Crown 8vo. pp. xiii + 245. With 10 plates and 18 text-figures.
Price 5i. net

"Nowadays," says Dr Lock in his Prt'/(7fc, "rubber enters so intimately
into the daily life of almost everyone, that there will probably be few to
whom the romance of rubber entirely fails to make an appeal."

To make the book suitable to as wide a circle of readers as possible the
author has aimed at combining an accurate account of the scientific side ot

Hcvid Rubber and Tea

rubber planting with a certain amount of practical information for the
benefit of the prospective planter.

The chapter-headings are as follows : The History of the Use and
Cultivation of Rubber, The Botanical Sources of Rubber, The Physiology
of Latex Production, Tapping Experiments, Hevea Planting Operations,
Harvesting Operations, Factory Work on the Estate, The Pests and
Diseases of Hevea, The Cultivation of Species other than Hevea Brasi-
liensis. The Chemistry of India-Rubber, The Manufacture of Rubber
Goods.

13

FLIES AND THE PUBLIC HEALTH

Flies in relation to Disease. Non-Bloodsucking Flies. By G. S. Grahat,
Smith, M.D., University Lecturer in Hygiene, Cambridge.

Demy Svo. pp. xiv^-292. With ^4. plates and 32 text-figures.
Price loi. 6./. net

The first volume in The Cambridge Public Health Series deals with
a subject which has hitherto received too little attention.

" The work done up to the present " he says " has been mainly of a
preliminary character. It has, however, established certain very important
facts; that many of the non-biting flies found in houses walk over and feed
on decaying substances and
excreta of all kinds, and that
their larvae develop in them ;
that occasionallv disease-pro-
ducing bacteria may be pre-
sent in these excreta; that
flies can carry bacteria on
their limbs and bodies for
several hours, and internally
for several days; that for
some days they can infect
substances, including human
food materials, over which
they walk or defaecate,. and
on which they feed ; and
that their habits are such
that they constantly infect
food with the bacteria they
carry. Further, the epidemiological evidence suggests that, when suitable
conditions prevail, flies may be highly important factors in the spread or
certain infectious diseases

In this book an attempt has been made to collect the most important
and reliable information available on the subject, and to arrange it in such
a manner that all who are interested in its various aspects may be able to
ascertain the present extent of our knowledge.

In order to meet the requirements of various classes of readers, those
portions of the book which are devoted to matters of general interest and
importance are printed in large type, and in them, as far as practicable, the
use of technical terms has been avoided. The details of bacteriological
experiments and technical descriptions ot important insects have been
printed in smaller type, for the convenience of medical officers, bacterio-
logists and entomologists."

Photograph (side view) of an unfed

fly (x7)

14

PROJECTIVE GEOMETRY— BUILDING CONSTRUCTION

The Principles of Projedi've Geometry Applied to the Straight Line and
Conic. By J. L. S. Hatton, M.A.^ Principal of the East London
College (University of London).

Royal Svo. pp. x+366. Price loi. 6./. net

The book aims at giving a pupil who has already mastered the portions
of Euclid usually read, most of the pure geometry required for an honours
degree at Oxford, Cambridge, London, or Manchester. The subject has
been considered primarily from the projective point of view, but consider-
able trouble has been taken to deduce the more important metrical pro-
perties of conies from the projective theorems with which they are
related,

In these days, when the analytical method seems to hold the field in
mathematics, the author hopes that the book may encourage the student
to pay due attention to the methods of pure geometry, ten years' experience
as examiner in the university of London having shewn him the value of a
mastery of the principles of pure geometry.

"The student whose tastes lie in the direction of pure geometry will find here a
mass of excellent material upon which he can exercise his faculties." — Sihool ll'orUi

Architectural and Building Construction Plates. Part L Thirty draivlngs
covering an elementary course for architectural and building students.
By W. R. Jaggard, A.R.LB.A.

Size 20" X 13". Price 6s. net in portfolio, or in 6 separate parts, is. 3</. net each

These drawings, which form the first volume of The Cambridge Technical
Series (see p. 30), were prepared by the author as a basis for a series of
Lectures upon Building Construction, and are intended to cover an -ele-
mentary course of instruction in that subject. The examples have been
carefully chosen from work actually carried out under the direction of the
author, and will form a progressive series of drawings adapted to the
requirements of architectural and building students just entering upon their
professional studies.

In some cases, the plates will form a ready source of reference, both for
architect and builder, upon items of construction not readily found in
text books.

For teachers of building construction the plates, supplemented by
models, carefully made from the data given, will be found most useful
in imparting knowledge of what is, admittedly, a difficult subject.

The drawings illustrate Brickwork, Excavating, Masonry, Carpentry,
"Joinery, Roof Coverings, Plumbing, and Standard Steel and b'on Sections.

15 1—9

THEORIES OF EDUCATION

The Little Schools of Port-Royal. By H. C. Barnard, M.A., B.Litt.,
Sometimf senior Hulrne Scholar of Brasenose College, Oxford.
Demy 8vo. pp. x+264. With 4 illustrations. Price yj. 6d. net
In his Introduction the author claims that the educational work of
the Port-Royalists is worthy of study, both for its own sake and for the
importance of its subsequent influence in France and elsewhere. At the
same time it is true to say that the doctrines and practice of the Little
Schools are but little known not only in this country but even in France
itself. The reason is not far to seek. Port-Royal's chief importance has
been theological and literary and the value of the Port-Royalists' con-
tribution to education has been very generally overlooked. Moreover the
treatises in which their educational theories are expounded are, for the
most part, rare and not easily accessible ; again, these details are, to some
extent, scattered throughout " Memoires " and histories and letters where it
is not easy to find them and where they are often intermingled with ex-
traneous matter of every kind. But, for all this, a sympathetic study of
the methods used in the Little Schools cannot fail to be a source of
inspiration to teachers of the present day.

Vives : On Education. A tramlation of the De Tradendis Disciplinis of
Juan Luis I ives together with an introduction by Foster IFatson, D.Lit.,
Professor oj Education in the University College of JVales, Aberystwyth.
Crown Svo. pp. clviii-fjzS. With a portrait of Vives. Price 5J. net

This is the first English translation of Vives' main educational work
and though Professor Watson admits that the most likely answer to the
question " Was Vives a greater thinker on educational matters than
Erasmus ? " would be " We never heard of Vives and we are perfectly
familiar with the importance of Erasmus," he claims at the same time that
"there is ?i prima facie case for enquiry, whether Vives in his desire for the
social good, and all questions bearing upon that standpoint, has not treated
the subject of education so fruitfully as to be emphatically the greatest
European educational leader of the first half of the sixteenth century."

" It is a book," says The Athenaeum, " that should be read by every
teacher, if only to show how old are most of the modern suggestions for
the improvement of education."

Steps toivards Educational Reform. Some practical suggestions for improving
our national system. By C. IF. Bailey, M.A., Headmaster, Holt
Secondary School, Liverpool.

Crown Svo. pp. viii+112. Paper boards. Price u. net

Aims at showing how mistaken is the view that State organisation and
subsidy can be a panacea for all educational ills, that there is a distinct
danger in increasing bureaucratic control and that the chief need is more
light and life in the schools themselves.

16

Complete List

of
Books published since i 5 April 1 9 1 3

THEOLOGY

The Book of the Prophet Jeremiah together <vjith The Lamentations. Re-
vised /Version. Edited, with introduction and notes, h\ A. JV. Stream,
D.D. Fcap. ivo. pp. liv + 382. With 3 maps. Price 3s. net.

In The CambriJge Bible for Schools series. Large additions and modifications have been
made embodying the results of theological research during the 32 years which have
elapsed since the publication of the first edition.

The Gospel according to St Luke. The Greek Text. Edited, tuith intro-
duction and notes, for the use of schools, by IV. F. Eurnside, M.A.,
Headmaster of St Edmund'' s School, Canterbury. Fcap. 8vo. pp. xxxvi
+ 272. JFith 2 maps. Price 3$. net.

A companion volume to Sir Arthur Hort's St Mark. The notes are as brief as
possible and those upon grammar and language are in a strictly subordinate position,
though striking differences between classical and Hellenistic Greek are marked. The
meaning of the Gospel is drawn out in such a way as to appeal to those who have
made sufficient progress to read the New Testament in Greek.

Scripture Teaching in Secondary Schools : « Report of a Conference held

at Oxford 22-23 ^P'''^ I9'3- Second Tear. Edited by H. Cradock-

JVatson, M.A. Crown ^vo. pp. viii + q^.. Price is. 6d. net.

Contains papers read by Canon Scott Holland, Dr A. C. Headlam, H, Cradock-

Watson, the Headmaster of Harrow, Canon Masterman, Principal Selbie, Professor Peake,

H. S. Wood, N. P. Wood, together with a bibliography which, it is hoped, will be

found serviceable by busy schoolmasters and others.

The Fourfold Gospel : Section I, Intr'oduction. By Edwin A. Abbott, D.D.
Demy \ivo. pp. xvi + ijS. Price2s.6d.net. (Diatessarica. Part X.
Section L)

This volume, utilising the results of the previous volumes of the series in the
elucidation ot Gospel words, aims at the elucidation of Gospel thoughts, and after
discussing the order ot the tour Gospels, treats of the order and arrangement of the
subject-matter in each ot them. The author lays special stress on the value of St John
as a historical document.

Miscellanea E-vangelica (I). By the same author. Demy ivo. pp.
xii + 100. Price 2s. net.

The Song of Songs. Edited by IF. IF. Cannon. See p. 7.

Early Latin Hymnaries. By J. Mearns, M.A. See p. 6.

Boanerges, By Rendel Harris. See p. 7.

17

THEOLOGY— GREEK

Paganism and Christianity) in Egypt. By P. D. Scott -Moncrief, M.A.
See p. 6.

The History of the Islands of the Lerins. By A. C. Cooper- Maruiin, D.D.
See p. 8.

The Early History of thS Liturgy. By J. H. Srawlry, D.D. See p. 7.

The Interregnum. By R. A. P. Hill, B.A., M.D. See p. 6.

GREEK

Theocritus, Bion and Moschus. Translated into English Eerse hy A. S. IFay,
D.Lit. Fcap. .^to. pp. via + 158. Price 55. net.

"The talent and sympathetic insight of Dr Way have enabled him to produce a
translation of these three classics which, taken all round, surpasses every previous
attempt to render their spirit into English verse." — Engiis/i Re-vieiv

Salamis. In easy Attic Greek. JFith introduction, notes, and vocabulary,

by G. M. Edwards, M.A., Felloiv of Sidney Susse.v College, Cambridge.

JFith a map and 3 illustrations, pp. .xvi -\- 78. Price is. bd.

In the series of Cambr'utge Elementary Classics. The text of Herodotus has been

much shortened and simplified and is intended for students in their second year of

Greek reading. The re-written text is mainly Attic ; but a tew Ionic touches have

been retained for their literary value ; and there is abundance of useful idiom.

A Short Syntax of Ne'w Testament Greek. Second edition. By Rev.
H. P. F. Nunn, M.A., St Johns College, Cambridge, sometime lecturer
at St Aidans College, Birkenhead. Crown 2ivo. pp. .v;7 + 172. Price
2s. bd. net.

A Selection of Illustrative Passages to the Syntax of Ne<w Testament
Greek, by Rev. H. P. I'. Nunn, M.A. Fcap. 'ivo. 28 pp. Paper
covers. Price bd. net.

A supplement to A Short Syntax of Neiv Testament Greek and incorporated as an
Appendix in the second edition of that work.

A Greek Vocabulary for the use of schools. By T. Nicklin, M.A.,
Assistant Master at Rossall School. E.xtra fcap. ?ivo. pp. .V//+104.
Price 2s. bd. net.

This vocabulary is constructed on quite original lines for the special benefit of young
students of Greek and consists of four lists :

(i) A main vocabulary of 1300 words, each of which occurs more than
20 times in Euripides and Thucydides combined ; in this list the
300 commonest words are specially indicated.

(2) 270 words found more than 15 times in the Gospels and Acts.

(3) 150 words from the Anabasis, not included in (i).

(4) 400 common words from- Demosthenes., not included in (1).
The total vocabulary thus contains about 2000 words.

18

LATIN— ENGLISH LITERATURE

LATIN

Quantity) and Accent in the Pronunciation of Latin. By F. IF. JVataiuay.
See p. 3.

An Elementary) Latin Grammar. By Arthur Sloman, M.A., late Head-
master of Birkenhead School. Crown Si'o. pp. xii -\- 170. Price is. bd.

Follows in the main the recommendations of the Joint Committee on Grammatical
Terminology and gives all the Grammar necessary for students trom the senior forms
of Preparatory Schools up to the usual standard ot entrance examinations at Oxford,
Cambridge and other universities.

Prima Legenda : First Tear Latin Lessons. By 'J. IFhyte, AI.A.f Classical
Mistress, Blackheath High School for Girls. Fcap. ^vo. pp. viii + 64.
Price IS. ^d.

Provides material for a year's work tor pupils beginning Latin at the age ot 1 1 or
12, and, according to a reviewer in Education, is a "splendid first year's course" which
"will make the study of Latin a delight."

The exercises, Latin-English and English-Latin, are in the torm ot simple stories.
The grammar includes the declension of nouns and adjectives, and the indicative mood
of the first four conjugations of verbs. In the appendix are some simple Latin versions
of nursery rhymes, and at the end of the book a complete Latin-English vocabulary.

Csesar. Gallic War. Book L Edited.^ with introduction, notes and voca-
bulary, by E. S. Shuckburgh, Litt.D. Fcap. pp. xxxii -\- 1^2. With
21 illustrations, T^ plans and a map. Price \s. bd.

A new edition of which the chief features are the marking of vowels long by nature
in the text and the inclusion of illustrations taken from authentic sources.

ENGLISH LITERATURE

The Cambridge History of English Literature. Edited by Sir A. IV. Ward,
Litt.D., and A. R. Waller, M.A. Fol. X. The Age of Johnson.
See p. 9.

Collected Literary Essays, Classical and Modern. By A. W. Verrall,
Litt.D. See p. I.

Outlines of Victorian Literature. By Hugh Walker, LL.D., and Mrs Hugh

Walker. See p. 9.

Edmund Burke. Thoughts on the cause of the present discontents.
Edited by W. Murison, M.A., Senior English Master, Aberdeen Gram-
mar School. Fcap. Svo. pp. xxxviii + 1 64. Price 2s. bd. Pitt Press
Series.

The Introduction includes a brief sketch of Burke's life, a resume of public affairs
{1760-70), a critical discussion of Tie Present Discontents, an appendix containing
Goldsmith's epitaph and the Middlesex petition (1769) and a chronological table; the
Notes are preceded by an analysis of the argument, and a summary of the pamphlet.

19

ENGLISH LITERATURE— FRENCH

Alexander Pope. Essay on Man. Edited with introduction, analysis of
the poem^ and notes, hy A. Hamilton Thompson, M.A., F.S.A. Fcap.
ivo. pp.xl+%%. IVith a portrait. Price 2s. Pitt Press Series.

Intended to supply the student with a readable text ot one ot the chief masterpieces
of 1 8th century poetry and with a short commentary and notes, which may serve to
elucidate the poem without becoming a formidable addition to it.

Byron. Childe Harold's Pilgrimage. Edited by A. Hamilton Thompson,
M.A., F.S.A. With portrait, introduction, notes and summary. Fcap.
ivo. pp. xxii + 2S6. Price 2s. 6d. Pitt Press Series.

Intended to supply a commentary upon the numerous historical, literary and
topographical allusions in the poem. Explanations of the more involved passages are
given where necessary, and quotations from classical and foreign writers, in cases where
they throw light on the text, are translated in the notes.

The Granta Shakespeare. Edited by J. H. Lobban, M.A.
The Merchant of Venice, is.
A Midsummer-Night's Dream, is.

Each volume has a frontispiece, introduction, notes and glossaries.

Henry Fielding. Journal of a Voyage to Lisbon. Edited, tvith intro-
duction and notes, hy "J. H. Lobban, M.A. Extra fcap. Sua. pp. xvi
+ 116. Price I s. 4^.

In The English Literature for Schools series. The editor holds that by this Journal
Fielding demonstrates his claim to stand beside Chaucer, Shakespeare and Scott, not
only by right of genius, but by his serene courage and large-hearted humanity.

FRENCH

The Song of Roland. Translated into English Ferse by A. S. Way, D.Lit.
Pott 4.10. pp. xvi -\- 144. Price 45. net.

Erckmann-Chatrian. Le Blocus. Chapters I — XHL Edited, with intro-
duction, notes and vocabulary, by A. R. Ropes, M.A. Fcap. %vo. pp. xvi
4 218. IVith tnap and plan. Price \s. bd. Pitt Press Series.
^ An edition designed for the Local Examinations, 19 14.

Erckmann-Chatrian. Le Blocus. Edited, with introduction, notes- and
vocabulary, by A. R. Ropes, M.A. JFith an appendix by Cloudesley
Brereton. Fcap. ivo. pp. xvi + 374. Map and plan. Price 2s.
Pitt Press Series.
A re-issue at a reduced price.

Erckmann-Chatrian. Madame Therese. Edited, with introduction, notes
and vocabulary, by A. R. Ropes, M.A. Fcap. ivo. pp. xvi + 2^0.
Map. Price y. Pitt Press Series.
A re-issue with Vocabulary.

20

GERMAN— MISCELLANEOUS PHILOLOGY

GERMAN

Schiller, Die Braui von Messina. Edited, with introduction and notes, by
Karl Breul, Litt.D., Ph.D., Schroder Professor of German in the Uni-
versity of Cambridge. Fcap.%vo. /i/. cro + 279. Price \s. Pitt Press
Series.

"Professor Breul has performed his task with his wonted skill and conscientiousness.
The introduction is particularly full, and the serious student will find its perusal stimu-
lating and profitable. The notes are full and accurate. There can be no doubt that this
will long remain the standard edition of ' Die Braut von Messina'." — School If'orlJ

Hacklander. Der Zaave. Adapted from Ein Schloss in den Ardennen.
Edited by G. T. Ungoed, M.A. Large crown 2ivo. pp. viii + 62.
Price 2s.

Designed tor pupils who have completed their first course in German. The text is
simple in vocabulary and will not require more than one term for study. The book
includes phonetic transcriptions ot short passages and of difficult words, a brief sketch
of the author's career,. questions on the narrative, grammatical exercises and subjects
for free composition ; it is primarily intended for use on the direct method.

Deutsches Heft. A German Note-book. Arranged by TV. E. IVeber, M.A.,
sometime Modern Language Master at Blackheath School, Bridlington
Grammar School, and Trinity College, Glenalmond. Fcap. ^to. pp. viii
+ 128. Price IS. 6d.

A companion volume to the French Note-book, Cahier Franfais de notes di-verses by
the same author. It is intended to help students, whether in class or self-taught, to
compile their own grammar and vocabulary, collection of idioms, etc. in the course of
their reading. The notes are methodically arranged with suggestive types heading
each page, and there is a full index.

MISCELLANEOUS PHILOLOGT

Phonetic Spelling : A proposed Universal Alphabet for the rendering of
English, French, German and all other forms of Speech. By Sir Harry
yohnston, G.C.M.G., K.C.B., Hon. Sc.D. [Cantab.). Crown Svo.
pp. vi + 92. Price y. 6d. net.

Intended to serve two purposes : (i) to set forth an ideal alphabet to be used for the
transcription of all tongues about to adopt a phonetic system of spelling or being tran-
scribed for the first time by missionaries and explorers ; {2) to explain how this alphabet
may be applied to the rendering of English and other European tongues.

"On our part," says Tie Pioneer (the organ of the Simplified Speling Sosieti), "we
rejois in Sur Harry'z help in creaiting an enlietend public opinion, ever our cheef wurc.
A shaic ov the hand tu aul hu ar triing tu maic a hiewai thru the SIou ov Despond."

21

CLASSICAL LITERATURE— HISTORY

Dictionary of the Hausa Language. By C. H. Rohinson, D.D., sometime
Lecturer in H<iusii in the University of Cambridge, Honorarx Canon of
Ripon. Volume /, Hausa-English. Third edition, revised and enlarged.
Crown %vo. pp. xiv + 4.26. Price 12s. net.
Contains about 4000 entries additional to those which appeared in the 2nd edition.

A\GaUa-EngUsh English-Galla Dictionary. By E. C. Foot. See p. 10.

CLASSICAL HISTORY AND LITERATURE

Collected Studies in Greek and Latin Scholarship. By A. TF. Verrall,
Litt.D. See p. i.

The Municipalities of the Roman Empire. By J. S. Reid, Litt.D. See p. 3.

Political Parties in Athens during the Peloponnesian War. By L. JVhibley,
M.A., Fellow of Pembroke College, Cambridge. Second edition. Crown
%vo. pp. via + 142. Price y. 6d. net.
A re-issue with a change ot price.

HISTORY

Early Wars of Wessex. By Albany F. Major. See p. 5.

The Cambridge Medieval History, folume II, The Rise of the Saracens
and the Foundation of the IFestern Empire. See p. 4.

The Ottoman Empire, J80i—i913. By IFilliam Miller, M.A. See p. 5.

Sion College and Library. By E. H. Pearce, M.A. See p. 8.

Exercises and Problems in English History, 1485 — 1820. Chiefly from
original sources. Compiled by IV. J. R. Gibbs, B.A., Senior Assistant
Master, IVest Suffolk County School. Crown ivo. pp. vi+lj^.
Price 2s. 6d.

The aim of the book is to present a type of exercise based on original sources which
does not necessarily demand an answer in the form of an essay and yet does not sacrifice
the worth of the problem. Thus, a page and a half is quoted (Ex. 28) from More's
Utopia and the following questions appended :

Examine carefully this extract and give a short summary of the results of the
increase of sheep runs.

Why did the sheep runs increase ?
Again a short extract from a proclamation (Ex. 66) is followed by the questions:
Whose proclamation is this .' Give reasons for your choice. What did the
author intend to do and why >

Great Britain and Ireland, 1485 — 1910. A history for lower forms. By
y. E. Morris, D.Litt., Bedford Grammar School. Crown %vo. pp. viii
+ 300. With 45 illustrations and 15 maps and plans. Price 2s.

Reprinted from the complete work for the convenience of those teachers who prefer
to make the di\'ision at 14S5 rather than at 1603.

22

LAW— MANUALS— ARCHITECTURE— MATHEMATICS

LAW

Outlines of Criminal Laiu. Fifth Edition, Revised, embodying the Forgery
Act, 1913. Based on Lectures delivered in the University of Cambridge.
By Courtney Stanhope Kenny, LL.D., F.B.A., Downing Professor of the
Laws of England. Demy ^vo. pp. xxx -\- '^ifi. Price los.

Papers set in the Special Examination in Laiv in the University) of
Cambridge, 1907— i9Ji. Crown 8vo. pp. iv + 80. Price 2s. 6d. net.

CAMBRIDGE MANUALS

Under the general editorship of P. Giles, Litt.D., and A. C. Sezvard,
F.R.S. IS. net in cloth, is. bd. net in leather.

71

72

73

74'

75
76,

77
78

79
80

FOLUMES 71—80

Natural Sources of Energy. By Prof. A. H. Gibson, D.Sc.

The Fertility of the Soil. By E. J. Russell, D.Sc.

The Life Story of Trisects. By Prof. G. H. Carpenter

The Flea. By Harold Russell '

Pearls. By Prof. JV. J. Dakin

Naval Warfare. By J. R. Thursfield, M.A. IVith an Introduction

by Rear-Admiral Sir Charles L. Ottley
The Beautiful. By Fernon Lee
The Peoples of India. By J. D. Anderson, M.A.
The Evolution of New Japan. By Prof. J. H. Longford
A Grammar of English Heraldry. By IF. H. St J. Hope, Litt.D.

ARCHITECTURE

Architectural and Building Construction Plates. By IF. R. Jaggard.

In The Cambridge Technical Series. See pp. 15 and 30.

MATHEMATICS AND PHYSICS

The Principles of Projective Geometry Applied to the Straight Line and
Conic. By J. L. S. Hatton, M.A. See p. 15.

Elementary Algebra. By C. Godfrey, M.F.O., M.A., and A. IF. Siddons,
M.A. Large crown %vo. Complete in one volume, with answers 4.S. 6d.
without answers 4.(. Or in two volumes at 2s. bd. or 2s. each. Fol.'-T
in two parts at is. and 2s. or is. bd. each.

23

MATHEMATICS AND PHYSICS

The Lams of Thermodynamics. By IV. H. Macau/ay, M.J., Fellow of
King's Collfgi; Cambridge. Demy 8vo. pp. viii + ~2. Paper covers.
Price 35. net. Cambridge Engineering Tracts No. 2.

The aim of this tract is to provide a connected and accurate account of the funda-
mental principles of thermodynamics, combined with a sketch of methods of applying
the theory in special cases, to supplement technical books on the subject.

Papers set in the Mathematical Tripos, Part I, in the University of
Cambridge 1908 — i9t2. Crown 8vo. pp. iv + 70. Price 2s. bd. net.

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24

MEDICINE AND BIOLOGY

Proceedings of the Fifth International Congress of Mathematicians,
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25

BIOLOGY— GEOGRAPHY AND TRAVEL

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26

GEOGRAPHY— ARCHAEOLOGY— EDUCATION

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27

BIBLIOGRAPHY— CAMBRIDGE UNIVERSITY

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28

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29

Announcements

THE CAMBRIDGE PSYCHOLOGICAL LIBRARY

Under the General Editorship of Dr C. S. Myers, University Lecturer
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Psychology. By Prof. Jamn Ward, Sc.D., F.B.A. [In the press

The Nervous System. By Prof. C. S. Sherrington, M.D., F.R.S.

The Structure of the Nervous System and the Sense Organs. By Prof.
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Prolegomena to Psychology. By Prof G. Dawes Hicks, Litt.D.

Psychology in Relation to Theory of Knoiv.edge. By Prof. G. F. Stout.

Mental Measurement. By IF. Brown, D.Sc.

The Psychology of Mental Differences. By C. Burt, M.A.

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3°

ANNOUNCEMENTS

have been selected are men of wide experience in the work of Technical
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Automobile Engineering. By A. Graham Clark, M.I.A.E., A.M.l.Mech.E.

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Applied Mechanics. By Etvart 5. Andmvs, B.Sc

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Physics for Engineers. By J. P. Torke.

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31

ANNOUNCEMENTS

PROriNCUL GEOGRAPHIES OF INDIA

Sir T. H. Holland, the general editor of this new series, writes in his
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Naval and Military Essays read at the International Historical Congress
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32

ANNOUNCEMENTS

FORTHCOMING BOOKS

The following books are also in the press and will shortly be ready:
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On the Art of Writing: Lectures delivered before the Uni'versity of
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The Literary Relations of England and Germany in the ' 17th Century.
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A Primer of English Literature. By JV. T. Toung, M.A.

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A Book of English Prose. 2 volumes. Edited by Percy Lubbock, M.A.

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Greek Art in relation to Modern Sculpture. (Two lectures.) JVith illus-
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The Reign of Henry V. J'ol. /, 1413— 1415. By J. H. JVylie, M.A.,
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The Puritans in PoTver. By G. B. Tatham, M.A.

Bartolus of Sassoferrato. By C. S. N. Woolf, M.A.

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The Parables of the Gospels. By L. E. Browne, M.A.

Elements of Neiu Testament Greek. By Rev. H. P. J'. Nunn, M.A.

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A National System of Education. By J. Howard IFhitehouse, M.P.

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Genera of British Plants. By H. G. Carter, M.B., Cn.B.

Merionethshire. By A. Morris.

Northumberland. By S. R. Hasclhurst.

33

* •:• '^ il^ ^ V ^

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Robert K. Nabours. Studies of Inheritance and Evolution in

Ortliopteia I. (With Plate VI, 3 Text Figures, and 2 Diagrams) 141

C. W. Richardson. A Preliminary Note on the Genetics of Fragaria.

(With Plate VII and 4 Text Figures) 171

W. E. Agar. Parthenogenetic and Sexual Reproduction in Sivto-

cephalus vetulus and other Cladocera 179

E. S. Salmon. On the Appearance of Sterile " Dwarfs " in Humulus

Liqndus L. (With Plates VIII and IX) . ... 195

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P. G. Bailey. Primary and Secondary Reduplication Series. (With

1 Text Figure) 221

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JOURNAL OF GENETICS

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Volume III APEIL, 1914 No. 4

(1,

ON HYBRIDS BETWEEN MOTHS OF THE GEO-

METRID SUB-FAMILY BISTONINAE, WITH (^^Tvcrk
AN ACCOUNT OF THE BEHAVIOUR OF THE «'".av,c^*
CHROMOSOMES IN GAMETOGENESIS IN LYCIA
{BISTON) HIRTARIA, ITJIYSIA (NYSSIA)
ZONARIA AND IN THEIR HYBRIDS.

By J. W. H. HARRISON, B.Sc, and L. DONCASTER, Sc.D.

PART I. GENERAL ACCOUNT OF HYBRID BISTONINAE.

By J. W. H. Harrison, B.Sc. ,

In the Entomologist for July 1910 I indicated that I had succeeded
in producing a number of new hybrids in the trio of genera included in
what may be called, for the sake of convenience, the Biston group. I did
not then, however, publish any details, either biological or otherwise,
concerning them. Since then, in the case of many of these hybrids, I
have worked out their detailed life history and have therein compared
them, point by point, with the parent species. Further, I have secured
several new and imj)ortant pairings ; not only is this so, but in the case
of most of them I have published an exhaustive account in Oberthiir's
Lepidopterologie comparee, Fascicule vii. pages 333 — 655. In case this
work is not readily accessible to English readers, I am writing this
summary of my results to accompany Dr Doncaster's statement of his
investigations into the oogenesis and spermatogenesis of Lycia hirtaria
and Ithysia zonaria, and their two crosses, hybrids denhami and
harrisoni. (Plate XVII.)

The hybrids reared may be divided into two sections.

(a) The Primary hybrids, i.e. those with pure species as parents.

(6) The Secondary hybrids, i.e. those in which either of the two
parents was a primary hybrid.

Journ. of Gen. iii 16

•230 Ilyhiifh of Bistonine Moths

The piiiiiaiy liybrids reared up to the present are as follows:

Lycia hirtaria ^ x Ithysia zonaria % (PI. XVII) = Lycia hybr. denliami.
Lycia hirtaria ^ x Ithysia go-aecaria % = L. Iiyhr. buluveci.

Lycia hirtaria (^ x Poecilopsis jwrnonariu J — L. liyhr. pilzii.
Ithysia zonaria. ^ x Lycia hirtaria % (P1.XVII)= X. /(j/tr. harrisoni.
Ithysia zonaria ^ x Poecilopsis lapponaria ? = /. hybr. nierana.
Ithysia zonaria ^ x Poecilopsis pomonaria ^ = I. hybr. lanc/ei.
Poecilopsis pomonaria ^ x Lycia hirtaria J = P. hybr. hunii.
Poecilopsis ponionaria J" x Ithysia zunaria. % = P. hybr. helenae.
Poecilopsis lapponaria ^ x Ithysia zonaria % = P. hybr. sniallmani.

Similarly, the secondary hybrids are appended :

Poecilopsis ponionaria ^ y. L. hybr. pilzii % = P. Iiybr. brouksi.
Lycia hybr. pilzii ^ x L. Irirtaria $ = L. hybr. burrowsi.

P. hybr. hunii ^ x L. Iiirtaria % = P. hybr. hnlli.

Up to the present, only one tertiary hybrid, that is to say a hybrid
with one of my secondary hybrids as a parent, has been obtained. This
form has been successfully reared to the pupal state and I trust to have
the pleasure of seeing the imagines next spring. Its parentage is
L. hybr. burrowsi ^ x L. hirtaria % and I am calling it L. hybr. adkini
in honour of my friend Mr. R. Adkin, who has assisted me in many ways
during the course of my many experiments.

I have no intention of giving a prolonged account of the comments
I made in my work in Lepidopteroloyie comparee upon the various
phenomena observed which demanded special treatment. Two of these
features, however (in my eyes at least), are so important that I have
reserved them for a special pajjer to be published shortly.

The various points are given below and a brief resume of the
discussion in my longer paper is added to each.

(1) The great constitutional strength of the larvae.

(2) The growing sterility of the printary hybrids as the specific
divergence between the parents increases.

I pointed out that " strength " in the case of all of the crosses
behaved as a Mendelian dominant, and that, granting the jjossibility of
aberrations (mutations) po.ssessing great constitutional strength, these
two factors alone would result, in some cases, in what could only be
classed as new species.

JOURNAL OF GENETICS, VOL. 3. N? 4.

PLATE XVII.

Ithysia (Nyssia) zonaria

Lycia (Biston) hirtaria

L, hirtaria 9x1 zonaria <?

I zonaria 9 x L hirtaria cf

Cambridge UniversiLy Press

J. W. H. Harrison and L. Doncaster -231

(3) Dominance of characters derived from the undoubtedly weaker
Ithi/siae.

This was treated at length and shown to depend on the fact that,
highly specialised as the Ithysiae are, they are yet, in many points, the
most primitive genus of the group in spite of the fact that, for the most
part, the genus Lycia is nearer the common phylogenetic ancestor of
both genera. In other words, we have a recurrence of common ancestral
features.

(4) The superior influence of the mule.

This is an experimental result that. I have noted in practically all of
the hybrids I have reared, not only in this family, but amongst the
Ennomids and Larentiads likewise. No reasonable explanation of this
fact has ever been suggested.

(5) The refusal of the food plants of the Ithysiae.

I have pointed out that this probably depends on the fact that, in
general, the Boarmiads are an arboreal group and that the Ithysiae
have almost certainly abandoned the family habit ; there is thus latent
in them a tendency to eat the same food jjlants as the Poecilopsis-Lycia
fraternity.

(6) The abnormal sex proportioJis yielded.

Four of the hybrids,

Lycia hybr. denhami,
L. hybr. bidoveci,
P. hybr. smallviani,
P. hybr. helenae,

exist only in the male sex under normal conditions.

It must not be supposed that this failure to give females is restricted
to this compact group of genera, for I have observed it in the case of
two other crossings I have made amongst the Boarminae, viz. :

(1) Epione paraUelaria ^ x E. apiciaria ? = hybr. isabellae

and (2) Tejjhrosia, crepuscularia c^ x T. bistortata % = hybr. bacoti.

Strong inbreeding of the parents producing two of the above hybrids,
on one occasion, resulted in the production of odd females, but the same
procedure was without effect whenever Lycia hirtaria took part in the

cross.

16—2

232 Hyhfids of BiMouiue Moths

Five of the hybrids,

L. Iii/br. jyilzii,
P. hyhr. huim,
L. hyhv. burruwsi,
P. hyhr. hulli,
L. hyhr. adkini,

yield the two sexes in ajjproximately equal numbers.
Three hybrids,

P. hyhr. langei,
I. hyhr. harrisoni,
P. hyhr. merana,

give a large excess of females.

Lastly, the sole form reared from a hybi'id $ , i.e. hybr. brovkai, gives
specimens which are hopelessly gynandromorphic, wings, body, genitalia,
antennae being built up of parts chosen at random, as it were, from
both sexes of the parents and gi-and-parents. It is worthy of note that
the larger and more robust the specimen, the more the male characters
predominate and vice versa.

(7) The tendency for the hybrich to emerge long before the parent
species.

In most of the hybrids, it appeared that the emergence was hastened
by a period varying from a fortnight to three months. Most curiously,
this varied with the sex for, in most cases, the acceleration noted in the
case of the females was very gi-eat compared with the slight displace-
ment seen in the males, although the divergence was never so great as
that observed in hybrid rohsoid = Larentia (Oporahia) dilututa ^ x Lar.
(0.) autumnuta $ . In this case, the females appeared about four and a
half months before the males and a similar period before both sexes of
the reciprocal cross riingei = L. (0.) autumnatu ,/ x L. (0.) dilvtata %.

It is not my intention to give here my thoughts on the import of
this, nor what appears to me to be the significance of the sex proportions
detailed above. I hope to be in a j^osition to publish my paper (already
planned out) before long.

(8) The wing development of the hybrid females.

The hybrid females, when produced, vary greatly in their wing
development ; those between hirtaria, fully winged, of course, in both
sexes, and one of the so-called apterous species, possess wings varying
enormously even within the limits of the same brood. This is a distinct

J. W. H. Harrison and L. Doncastbr 233

consequence of the fact that, whilst there is but little difference in the
wings of various specimens of hirturia females, in the apterous forms
there is an extraordinary amount of variability.

In lapponaria, for instance, the wings vary from mere vestiges to
long lanceolate appendages and similarly with pomonaria; zonaria on
the contrary, although not quite constant, for it varies slightly in the
same direction, is almost so. The hybrids have thus to combine the
immutability of the female wings of hirturia with the varying forms
developed in the other species. The result is that, whilst in most cases
we have wings produced resembling very roughly ordinary male wings,
although only two-thirds of the expanse, in many, we have long, narrow
pointed wings, and in others, wings most curiously shortened, giving one
the notion tliat a piece has been cut fi-om them. As one might expect,
the variation is least in the zonaria ^ — hirtaria % cross and greatest in
the two hirtaria — pomonaria crosses.

Nor is the variation confined to size and shape for, strange to say,
the wing scales vary simultaneously ; the smaller the wings the greater
the tendency for the .scales and fringes to become bristle or hair-like as
in normal apterous females. Further, although the rudimentary wings
of the apterous forms are not melanic, nevertheless the smaller the
wings in the hybrids the gi'eater the melanic tendency, which, however,
it must be remarked, is present in all of the female hybrids.

In crosses between the apterous forms, in the majority of cases, the
wings are small and constant, and seem greatly influenced by the wing
form of zonaria.

The tendency of hybrid brooksi to produce forms with all the
possible wing sliapes combined in one specimen has already been
noticed.

The wing forms of the other two secondary hybrids demand special
attention. The females yielded by the crossing hunii ^ and hirtaria %
possess forewings about five-sixths of the normal wing expanse of
hirtaria J* but with the inner margin shortened, and the costa and
termen strongly rounded. The hindwings, except that they are pro-
portionately stouter and broader, are not unlike those of hunii $ . The
scaling of both sets is that of hirtaria % ; the markings, too, except that
the fringes are quite black, follow hirtaria.

The hybrid between pz'kii ^ and hirtaria $ very curiously produces
females, which, except for the slightly rounded costa and termen, have
both sets of wings almost, but not quite, as fully developed as those of
hirtaria $ . If the predominant influence of the male, as mentioned

234 Hi/bn'ds of Bisfonine Moths

above, is carried thnnigh both generations this would explain the
difference in form between two hybrids so nearly the same in blood,
both being three i|narters hirturia and one quarter /)o;»o?)ffn'((.

PART II. ON THE CHROMOSOMES IN GAMETOGENESIS
OF THE MOTHS LYCIA {BISTON) HIRTARIA AND
ITHYSIA {NYSSIA) ZONARIA, AND IN THEIR HYBRIDS.

By L. DONCASTEE, Sc.D.,
Fellow of Kinc/'s College, Ciimhridge.

The work of which this paper gives a preliminary account was
undertaken to find out, if jjossible, any eytological cause for the fact
that reciprocal crosses of Lycia (Biston) hirtaria and Ithysia {Nyssia)
zonaria give different results as i-egards the sexes of the offspring.
Mr J. W. H. Harrison, who very kindly supplied me with the hybrid
material used, has described the results of the two crosses, together with
other hybrids, between species of the same genera. Ftir the present
purpose the important tacts ai-e that the ci'oss zonaria J x Iiirtaria </
produces only male offspring, resembling zonaria somewhat more nearly
than hirtaria; hirtaria % x zonaria ^ gives both sexes, with a con-
spicuous excess of females (more than 2 $$:!(/'). The males of this
second cross are not strikingly different from those of the converse cross ;
the females are remarkable in having small flightless wings. (Plate
XVII.) The wings of the hirtaria $ are of normal size and are capable
of flight; those of ^o«((/'('(f are vestigial, so that in this respect the h3'brid
is intermediate '.

Although the observations which I have been able to maki' on the
small supply of material hitherto available have not led to very satis-
factory results with regard to cause of the difference in sex-determinalion
in the reciprocal crosses, the general behaviour of the chromosomes in
the gametogenesis of the hybrids is so remarkable that I think it worth
while to publish an account of it at once. The chromosomes of the two
species are vei-y different from <ine another, and in the hybrids most of
the chromosomes derived from each parent can be recognised at a
glance.

' It should be noted that the reduction of the wings, wliich is a secondary sexual
character of zmmrta, is transmitted by tlie zonaria male.

J. W. H. Harrison and L. Doncaster 235

Further, in the maturation divisions, as was found by Federley in
hybrids of species of Pyguera}, most of the chromosomes are unpaired
and divided singly, so that the spermatocyte divisions have nearly as
many as the spermatogonia and oogonia, but my results differ from
those of Federley in the fact that a few of the kirtaria chromosomes
pair with some derived fi-om the zonaria parent, and that there is a
distinct S3mapsis stage in the prophases of the maturation divisions.
The methods of preservation, etc., employed were the same as those
described in my papers on gametogenesis of Abraxas, with the exception
that the gonads were dissected out in water, which I have found, since
those papers were written, to give better results than Ringer's fluid.
The material of the pure species was obtained from quite young pupae
purchased fiom Mr L. W. Newman ; that of the hybrids, from larvae
and pupae given me by Mr Harrison.

1. The chromosomes of i. (B.) Iiirturia.

(a) Oogenesis. I have very few satisfactory figures of the oogonial
divisions, but there seems to be little doubt that the chromosome number
is 28. Plate XVIII, fig. 2 shows an oogonial equatorial plate, with 28
chromosomes varying considerably in size, and including six which are
noticeably smaller than the remaindei'. I have one figure in which in
addition to a similar group <jf 28, there is a body just outside the circle
which might be a chromosome. I am inclined to believe that it is not
a chromosome, but an extra-nuclear body (fig. 1).

A typical synapsis stage succeeds the last oogonial divisions, followed
by a growth phase of the oocytes in which a relatively small number of
fairly thick threads form a spireme. As the oocytes enlarge, these
threads shorten in the usual way, and become faintly double, and by
the time they have become elongated rods about thirteen can be counted
(fig. 3). They then contract still fiu'ther t(j double, or often faintly
quadruple, bodies, of which again about thirteen may be counted.
Meanwhile a chromatin-nucleolus has been conspicuous, and still remains
much more sharply defined than the other chromosomes. It is com-
poimd, and in the many figures is seen to be composed of four parts, two
small and two larger. The latter are fi'equently seen to be unequal in
size. If the chromatin-nucleolus may be regarded, as in other forms, as
a " sex-chromosome " there is thus perhaps an indication of its being
unequally paired. There is no regularity, however, in its appearance ;
the parts may be together or widely separated, and are sometimes seen

' H. Federley, Zeitsclir. f. indiikt. Ahstamm. tiiiil Veierbiuiijdekre, ix. 1913, p. 1.

236 Hjjhr'nh of B!>itonme Motha

themselves to be composed of smaller units which show various degrees
of separation. The counting of the double rods is not quite easy;
usually thirteen seem to be present, but in some cases I cannot find
more than twelve. If, as seems probable, there are thirteen, the larger
and smaller members of the chromosome-nucleolus cannot consist of a
large and small pair of united chrcimosomes, as occurs in the spermato-
genesis (see below). Seiler has described the " sex-chromosome " in the
female of the moth Phragmatohia fidiginosa as being double', and the
doubleness of the chromatin-nucleolus seen in hirtaria may perhaps be
due to the same cause. As I have not investigated the polar divisions
of the eggs, I cannot test the truth of this suggestion.

(b) Spermatogenesis. I have a number of clear figures of the
spermatogonial divisions, and as in those of the oogonia there are clearlj'
28 chromosomes, of which si.K are smaller than the rest, and four of these
conspicuously smaller (fig. 4). In the gi-owth-stages of the spermatocytes
a typical sjmapsis of the spireme occurs. As in other Lepidoptera, the
stages are less easy to arrange in order than in the ovaries, but there is
no doubt that a thin spireme thread, closely withdrawn to one side of
the nucleus, is succeeded by a thicker and looser spireme (figs. .5, 6),
and this by a reticulum, from which the double chromosomes of the first
spermatocyte division arise by a process of concentration.

In the younger spermatocytes there are usually two chromatin-
nucleoli; in the older ones commonly four, two larger and two small.
These may be united so as to give three or two, a large one commonly
being paired with a small one. I cannot find any constant dift'erence in
size between the two pairs ; one of the larger or of the smaller is some-
times of greater size than its fellow, but this does not seem to be
regular. The stages in which the chromatin threads contract into
chromosomes are so confused that I am unable to count them, or in
many cases even to distinguish the ciiromatin-nucleolus from the other
chromosomes.

The primary spermatocyte divisions are usually extremely clear and
diagiamraatic, but are somewhat difficult to interpret, for in place of the
14 chromosomes which one would expect they constantly have only 13.
Careful inspection, however, shows that one of the largest is comj)ound,
and consists of a large chromosome to which a very small one is attached
(figs. 7, 8). That the large one is double (i.e. composed of an ecjually
matched pair) is often quite clear, but the small one attached to it

' J. Seiler, Zool. Aiizeitjer, xli. 1913, p. 246.

-T. W. H. Harrison and L. Doncastrr -237

usually shows no sign of doubleness, and on the spindle when seen side-
ways it sometimes appears to be attached to one half only of the large
one, as if it were going entire to one pole of the spindle. This, however,
appears to be exceptional ; the compound chromosome is usually recog-
nisable only with difficulty in side view (fig. 9, a, b, c). If it were the
case that the small chromosome went undivided to one pole like a
heterochromosome, the second spermatocyte equatorial plates should
be of two kinds of equal frequency ; half of them should have thirteen,
and half either fourteen, or thirteen of which one should be compound.
All these conditions in fact are found, but the great majority have
thirteen of which one may be more or less clearly composed of a large
and small member attached to each other with varying degrees of close-
ness. In some, perhaps the majority, only thirteen apparently simple
chromosomes are visible ; in others a large one clearly has a small one
attached to it, and in a few fourteen separate chromosomes may be
counted (figs. 11, 12, 13). In one of the latter class of figures, the
largest chromosome has clear indications of being compound although
there are thirteen sej)arate chromosomes in addition to it, an arrange-
ment which is undoubtedly abnormal (fig. 14).

Usually there are two chromosomes, in addition to the small member
of the compound one, which are conspicuously smaller than the rest,
but, probably according to the depth of staining and the position of the
chromosomes on the spindle, there is some variation, so that only one,
or sometimes three small ones, may be visible. It is thus not easy to
identify any one chromosome with confidence, but when fourteen are
visible, one very small one is usually, if not always, in close proximity
to one of the largest. It is clear from this account that equatorial
plates of the second spermatocyte divisions do not make it quite certain
whether one chromosome goes over undivided in the first division or
not. Unfortunately I have found only one case of a first division
anaphase in which the chromosomes of both groups can be counted
(fig. 10), and this is not entirely unequivocal. In each group there are
thirteen chromosomes, one of which shows signs of being composed of a
larger and smaller unit in each group. This would indicate that the
double chromosome divides equally, a large and a small portion going
to each pole. In one group, however, there is a body outside the group,
and at a different level in the section, which might possibly be a
chromosome. It is nearer the pole than the group of thirteen, and I
am fairly confident that it is not a chromosome but an extra-nuclear
body. Such occur regularly in the spermatocyte cells.

238 Hi/bruls of Bistonine Moths

Although, therefore, there are some appearances whicli suggest the
presence of a lieterotropic chi'oinosnine in the mahi, I am inclined to
believe that these are deceptive, and that one of the smallest chromo-
some jjairs constantly unites with one of the largest in the first
spermatocyte division, and divides normally, but that the closeness of
the union varies in different cases in the second division, so that some-
times thirteen, sometimes fourteen, appear to be present.

2. The chroniosonies of Ni/ssia (Tthi/sid) zoiiaria.

(a) Spermatugenesis. A glance at a spennatogonial equatorial
plate shows at once that the chromosomes are very unlike those of
L. (B.) hirtaria. Instead of 28 rather large chromosomes, there are a
very large number of extremely small ones. I have found no figure in
which they can be counted with accuracy; they tend to come into
contact and one can oidy make nut with confidence that there are over
100 (fig. 15).

The synapsis and growtJi phases are closely similar to tliose of
hirtaria, except that the "bouquet" stage (fig. 16) is less typical; as
in that species there are sometimes four chromatin-nucleoli, consisting
of a larger and smaller })air, but more frequently tht'.se are united to
form three or two. They are very nearly of the .same size as those of
hirtiiria. The prophases of the first spermatocyte division are clearer
than in hirtaria owing to the much smaller size of the chromosomes,
and it can be seen that the chromatin-nucleoli approach one another
more and more closely, and finally unite into a single roundeil mass in
which no division can be seen.

The primary spermatocyte (li\isions show equatorial pl.iti's of really
surprising beauty and perfection (fig. 17). The chromosomes lie
absolutely in one plane, and wiiJely separated from one aiiothei', so that
there is not the smallest difficulty in counting -56 with complete
certainty. Of these, two are noticeably larger than the rest, and
u.sually two others intermediate in size. The number 56 is so clearly
and certainly shown, that one may conclude with confidence that the
spennatogonial (diploid) number is 112, that is to .say, four times that
of hirtaria (28). The largest chromosomes of zonaria are of about the
same size as, or possibly even smaller than, the smallest of hirtaria. In
metaphase and anaphase it is seen that each chromosome is dividing in
the normal " heterotypc " manner, the diverging halves being connected
by double strands, and I can find no evidence of unequal division, nor
of the union of two pairs, such as occurs in hirtaria.

J. W. H. Harrison and L. Doncaster 239

The second spermatocyte division resembles the first very closely,
except that the chromosomes are half the size ; 56 can easily be counted,
of which four are larger than the others (fig. 18).

Some follicles of both zonaria and hirtaria have abnormal spermato-
cyte divisions, leading to spermatozoa without nuclei, as in Pygaera,
Abraxas, etc.

(6) Oogenesis. My zonaria pupae were too old to give satisfactory
observations on the oogenesis. The ovaries were already large, with
eggs at the lower end of the tubes in which a considerable amount of
yolk had been deposited, so that the tubes were becoming moniliform.
I have found no diploid mitotic figures in which the chi-omosomes can
be counted accurately. The younger oocytes at the top of the tube had
already undergone synapsis, and were in the stage with the chromosomes
aiTanged under the nuclear membrane. Completely accurate counts of
this stage are scarcely possible when the chromosomes are numerous ;
it can only be .said that there are between .50 and 60 small double
chromosomes, and a composite chromatin-nucleolus of which the two
largest portions are almost always of recognisably unequal size.

3. Chromosomes of the Hybrids.

(t() Zonaria % x hirtaria ^. This cross gives only male offspring.
My material consists of testes of two lai'vae shortly before pupation,
and of one pupa about three weeks old. The larval testes contain no
divisions later than the spermatogonia ; the pupal testis has also first
and second spermatocyte divisions, and contains spermatids in an
advanced stage of development towards spermatozoa.

The spermatogonia! equatorial plates show at a glance two kinds of
chromosomes — comparatively few large ones intermingled with a nnich
larger number of small ones.

A count of a very good figure (fig. 19) gives 5.5 — 57 small and 14
large ; although this cannot be i-egarded as absolutely accurate, the
error certainly does not amount to more than two or three small
chromosomes at most. The theoretical expectation is 70, so that it
may be assumed that complete iiaploid sets of hirtaria and zonaria
chromosomes are present.

The spermatogonial divisions are succeeded by a stage in which a
thin spireme is contracted to one side of the nucleus, after which
the thread thickens somewhat and becomes reticular, but the typical
"bouquet" stage (pachynema), which is found in pure hirtaria and less

240 Hyhrhh of Bhtonine Mothx

typically in zoiiuriu, seems not to occur (figs. 20, 21). In the mature
.spermatocyte there are either one or two chromatin-nucleoli which are
clearly compound ; the parts show less tendency to become separate
than in either of the pure species.

Federley, in his work on hybrids between species of Fygaera, found
no synapsis (synizesis) stage in the hybrids ; in the present case,
although most of the chromosomes fail to paii', so that there is almost
the diploid number in the spermatocyte divisions, there is no important
difference between the spermatocytes in the earliest growth stage of
the hybrids and those of the parent species. Tlie lact, however, that I
have found no " bouquet stage " with thick thread still contracted to
one side of the nucleus, is probably to be correlated with the fact that
most of the chromosomes fixil to pair. Federley's failure to find any
synapsis (synizesis) in Pygaera hybrids may possibly be due to his
material being too old. In the present case I find such a stage very
frequently in larval testes, but not in pupal testes about a month older,
in which the majority of the follicles contain advanced spermatocytes,
and the earlier stages are scarce. The various stages seem to (Overlap
less in the hybrids than in the pure species.

The spermatocyte division figures are veiy remarkable, and are not
conspicuously different from those of the spermatogonia. I have a
number of very perfect figures, and in all it is quite clear that the
chromosomes are nearly in the somatic number (figs. 22, 23), as was
found by Federley in his hybrids with Pygaera spp. Careful counts
.show, however, that the full somatic number is not present, and that
some pairing of chromosomes has taken place. It is not easy to draw
the line quite clearly between the large and small chromosomes, for as
was said above the larger zonaria are similar in size to some of the
smaller of hirtan'a. In the first spei'matocyte equatorial plate there are
always about 12 or 13 which ai-e certainly larger than all the rest, and
most of these may be regarded as hirtaria chromosomes. The number
of small ones is commonly about 50 ; careful counts have given 50, 50 or
51, and 51 or 52, in the three best figures I can find. Another fair
figure (slightly oblique, fig. 23) in which I cannot find that any chromo-
somes are omitted from the section, gives 13 large and about 40 small,
and others have given intermediate numbers ; it is possible that some
are covered by others, but it is unlikely that so many would disappear
in this way, and probably the smaller number may be due to the fact
that more chromosomes find mates in some cases than in others. There
seems no doubt that the total does not amount to the theoretical number

J. W. H. Harrison and L. Doncastbr 241

of 69 or 70 (56 + 13 or 14) which would be expected if no pairing of
chromosomes took place at all. This is further confirmed by the fact
that there are constantly 12 — ^14 large ones. Hirtaria spermatocytes
have eleven large and two small, and the largest of zonaria are hardly
big enough to be classed in the '■' large " group. If, however, the larger
of the zonaria chromosomes paired with the smaller of the hirtaria
chromosomes, these would make additional large ones in the hybi'id
spermatocyte equatorial plate. Several, in fact, often appear to be
double, and as the number in the best figures adds up to 65, it may be
concluded that about five zonaria chromosomes (probably the largest)
pair with five derived from hirtaria, while most of the rest remain
unpaired. An examination of the prophase figures of the first spermato-
cytes just before the nuclear membrane disappears, shows, in addition
to several clumps of three or more chromosomes together, a large
number of single chromosomes of various sizes, and among them a few
which are paired, either equally or unequally (fig. 24). Further con-
firmation of the evidence that some of the chromosomes are paired in
the first spermatocyte division and that the smaller number counted is
not due to error, is found in the tact that almost exactly the full
number can be counted in the spermatogonial divisions, which are
smaller and less easy to examine accurately than those of the spermato-
cytes. If it is easy to count very nearly 70 in the spermatogonia, it is
hardly possible that the smaller number in the spermatocytes can be
duo to error.

In the first spermatocyte division, it appears that not all the
chromosomes divide. Most undoubtedly do, but some show no signs of
division in metaphase when seen from the side of the spindle (fig. 25),
and the appearance of the secondary spermatocyte equatorial plates
confirms this supposition. The chromosomes in these second division
figures are less easy to count with complete accuracy, but it is not
difficult to get a fairly close estimate of their number, and this is almost
constantly less than in the first division (fig. 26). Counts have given
9 large and 42 small, 9 large and 46 small, 12 large and 42 small. In
one case, where I counted 12 large and 52 small, it is almost certain
that several had already divided, and that the halves were counted as
separate chromosomes.

{h) Hirtaria % x zonaria ^ . This cross gives a preponderance
of females, with some males. The ovaries of full-grown larvae are
extremely small and difficult to find, and fi-om the three female larvae
sent me by Mr Harrison I only succeeded in getting one ovary.

242 Hyhriih of Blxtoiune Moths

I allowed one larva to pupate ; it was fortunately a male and the testis
was preserved about two weeks after pupation.

The single larval ovary is most unfortunately not very well pre-
served. It contains only one oogonial equatorial plate which is
sufficiently in face for the chromosomes to be seen at all clearly, and it
is not good enough to provide an accurate count of the chromosomes.
It shows, however, a mixture of large and small chromosomes such as
I have described in the spermatogonia! plates of the converse cross.
Other odgonial divisi<ins, which lie more obli(|uely, also show chromo-
somes of two very different sizes. Owing to the defective preservation,
the development of the oocytes cannot be made out completely. There
is a stage with a thin spii'eine thread massed at one side as in the
normal early synapsis stage, and lower down the tube there are larger
nuclei with a thicker thread, but I have not found any nuclei in which
the thread has segmented into sejiarate chromosomes, such as are found
in normal older oocytes. The ovary, however, is so small that this
might perhaps not have occurred at this stage of development even if
normal synapsis had taken place. Since the development of the
chromosomes is much more clearly seen in oocytes than in spermato-
cytes of Lepidoptera, it is disappointing that in my only ovary the
stages required are not represented.

Spennatoffenesis. The testis of the single male pupa is large and
well develojjed, and contains all stages up to the spermatids, but no
nearly mature spermatozoa such as would probably occur in either of
the parent species at the same date. The spermatogonia! divisions are
like those of the converse cross ; I have not been able to make counts
which I can regard as completely accurate, but in the equatorial plate
from which fig. 27 is taken the chromosomes are clear enough to give
some approach to accuracy. I count in tliis plate 14 large and 55 small ;
in other plates I have counted only 11 or 12 large, and it is probable
that two or three reckoned as large in the group figured are zonaria
rather than hirtaria chromosomes. It is also possible that two or three
small ones counted as two are really single ones in division. The
number is sufficiently near to the theoretical expectation of 70 to make
it almost certain that complete sets from each parent are present.

The early stages of the development of the spermatocytes show
quite clearly cells with a fine spireme contracted to one side of the
nucleus ; it does not differ conspicuously from the corresponding
" contraction-phase " of synapsis in the pure species (fig. 28). After
this stage the nuclei enlarge considerably, and the thread takes the

J. W. H. Harrison and L. Doncaster -243

form of a reticulum, as in the converse cross (fig. 29). There are
usually at this stage two chromatin-nucleoli, each of which is double,
with halves which are slightly unequal ; the parts of one or both may
become separate, so as to produce three or four distinct masses.

The testis includes good equatorial plates of both first and second
spermatocyte divisions. Counts of first division figures have given
numbers ranging from .52 to 60 chromosomes (figs. 30, 31); in those
with the higher numbers it is possible that one or two dividing pairs
have been counted as two singles in each case. There seems no doubt
that the total number is not more than 60, and usually several less, and
as in .some figures as many as 18 have been reckoned as large, it is
practically certain that a considerable amount of pairing has taken
place, since only about eleven large ones are introduced from the
hirtarai parent, and the remaining large ones must be formed of pairs
of small ones. This is confirmed by the second spermatocyte divisions,
in which only about 10 large ones are counted; the small chromosomes
in this division are so small as to be counted with difficulty, but the
total of second equatorial plates is clearly not far from 50, made up of
about 10 large and 40 small (fig. 32). It is probable, therefore, that
in the cross hirtaria $ x zonaria ^ more chromosomes find mates in
the maturation divisions than in the converse cross, in which over
60 are usually present in the first spermatocyte divisions, but as a
lower number was found in some equatorial plates of this cross, it is
probable that the number which pair varies somewhat in different cells.

Condumjns and Diaciinsioii.

The general conclusions which may be drawn from the observations
described are then as follows: (1) there are 28 somatic chromosomes in
hirtaria, giving 13, of which one consists of a large united to a small, in
the spermatocytes. (2) There are 112 somatic and 56 spermatocyte
chromosomes in zonaria which are much smaller than most of those of
hirtaria. (3) The spermatogonia! number of the hybrids, whichever
way the cross is made, is as nearly as can be counted 70, of which
fourteen are conspicuously larger, and of these twelve are derived from
the hirtaria parent, the other two from zonaria. (4) The early synapsis
stage of the spireme in the young spermatocytes and oocytes of the
hybrids does not differ greatly from that of the pure species, but it is
apparently not followed bj' a normal pachytene ("bouquet") stage with
thick coiled thread. (5) Comparatively few of the chromosomes are

244 Hybri(h of Blstonlne Moths

paired in the spermeatocyte divisions, so that these have only from 5 to
15 fewer than the somatic number. It is (luitc clear, however, that
some chromosomes, the number of which is small and probably some-
what variable, do find mates in the maturation divisions, and that there
are probably homologous chromosomes in the two species which pair
together, while the remainder are unable to do so.

I do not projjose to discuss at length the observations of which only
a condensed account has been given, firstly because to a considerable
extent they are similar to those of Federley, and he has discussed the
questions raised at considerable length ; and secondly because I hope
next year to complete the investigation on additional material of the
same and other species, which should make a fuller understanding of
the problems jjossible. There are, however, a few jMints which seem to
require comment.

The chief points of interest are (1) the diffei-ence in chromosome
number in nearly related species, and the behaviour of the chromosomes
in the hybrids, and (2) the possible causes of the fact that the cross
z&naria $ x hirtaria </" gives only male offspring, while the converse
cross gives both sexes with excess of females.

In the former group of phenomena we have important data on the
vexed question of the individuality of the chromosomes. Since there
are exactly four times as many chromosomes in zonaria as in hirtaria,
and since the zonaria chromosomes are so much smaller that the total
amount of chromatin, as judged by the size of the resting nuclei, is
approximately equal in the two species, it seems a fair inference to
suppose that the hirtaria chromosomes niay be regarded as compound,
made up of units corresponding with the separate chromosomes of
zonaria. Since however there are large and small chromosomes in
hirtaria, and some in zonaria which are noticeably larger than the rest,
it is probable that not every hirtaria chromosome is made up of four
^jarts each of which corresponds with one zonaria chromosome, but that
the larger hirtaria chromosomes are made up of more than four units,
and the smallest ones are probably single. That is to say, if there are
56 units in the haploid gmup of zonaria, there are the same number of
units in hirtaria, but these are combined with one another in various
ways so that only 14 (or 13) chromosomes appear. For example, if the
56 haploid zonaria chromosomes are taken as units, the haploid set of
hirtaria might be made up of four chromosomes each coiiiposed of six
units, five of five, one of four, and three of one unit each. The total 56
is thus made uji (4 x 6 + 5 x 5 -t- 1 x 4 + 3 x 1). This is of course not

J. W. H. Harrlson and L. Doncastbr 245

the only possible arrangenient ; I give it merely as an illustration of
the way in which I regard the hirtarki chnunosomes as perhaps being
composed of a variable number of units each corresponding with one
zoniirid chromosome. It is of course probable, if the hirtaria chromo-
somes are compound, that those oi zonaria are so also, and that the true
" unit," whatever that may be, is smaller than either. Meek' has shown
that the chromosome lengths in various species of insects and other
animals are terms of an arithmetical series, as if they were composed of
units of standard length, and the comparison nf hirtiiria with zonariit
leads by a different route tn a somewhat similar conclusion.

Another aspect of the individuality question is touched by the
phenomena of the maturation divisions of the hybrids. I have shown
that although the majority of the chromosomes fail to pair in synapsis,
it is certain that some of th(un do so. The number of those which pair
is sometimes not more than ten (five pairs), giving a maximum number
of 65 in first sjDermatocyte equatorial plates in the cross zonaria % x
hirtaria ^ ; in other cases, especially in the converse cross, the number
of pairs appears sometimes at least to be larger. If I am right in
suggesting that about three of the small hirtaria chromosomes corre-
spond with single zonaria units, we might expect these to pair normally
together, and the indications of occasional unequal pairing suggest that
some of the compound hirftiria chromosomes may pair also, perhaps le.ss
regularly, with zonaria units. This would lead to the conclusion, not
that chromosomes are individuals in the sense of being indivisible units,
but that they are composed of units, and that pairing in synapsis i.s due
to some affinity between chromosomes made up of similar components.
This is of cour.se the hypothesis which has been widely accepted on
evidence of a different kind, in connexion with the hypothesis that the
chromosomes are the bearers of Mendelian unit characters.

If there is any truth in this view, it may give some clue to the
baffling problem of interspecific sterility. The hirtaria — zonaria hybrids
are completely sterile. If the haploid set of 56 chromosomes of zonaria
consists on the whole of the same units as the 14 haploid chromosomes
of hirtaria, combined in different ways, the sterility between the two
nearly allied species may have arisen simply from a difference of
grouping. Since the units are gi'ouped differently, the chromosomes
cannot pair properly in synapsis, and this may be the cause of the

' C. F. U. Meek, "A metrical analysis of chromosome complexes, etc.," Phil. Trans.
Roy. Hoc. Vol. B. 203, p. 1, 1912.

Jouru. of Gen. in 17

246 Hjihrid^ of Bi.'ifrmiur Mot]i<<

sterility. It is true that Federley's Pyguera hybrids were not sterile,
but that may be because little or no pairing took place, so that the
resulting gametes would be less abnormal than in a case where some
chromosomes pair and others do not.

The second point mentioned at the beginning of this discussion wa.s
the one which led in the first instance to my undertaking the work —
the possible cause of the fact that only males are produced from the
cross sonaria % x hiiiuria ^. Similar results have been obtained
with other species by various ob.servers, notably the cross Tephrasin
histortafa $ x 2\ crepusctilariti ^ described by Tutt'. Brake and
Goldschmidt's observations on Lijinaidna dlspur and L. juponica may
also be compared, in which they find that dispar % x juponica J" gives
males and gynandromorphs, although the converse cross gives normal
males and females". As Mr Harrison has shown, true hermaphrodites
occur in certain crosses with the genus Biston, but not with those used
in the present case.

My observations on the chromosomes do not yet give any conclusive
results with regard to the cause of these sex-phenomena, but there are
certain indications which perhaps deserve mention. It is probable that
in Lepidoptera there are two similar " se.x-chromosomes " in the male,
and that one of these differs from the other, and may be regarded as
lacking the male determiner, in the female. This has been suggested
on the ground of the facts of sex-limited inheritance (Abruxan); the
suggestion is supported, though not proved, by the observations of
Seller, of which only a preliminary account has been published^; and
I have obtained additional evidence for it from a study of the chromo-
somes in a strain of Abru.rus which in each generation produces families
consisting only of females. In a first account of this work* I have
shown that females of this strain have 55 instead of 5t) chromosomes,
and further work, an account of which I hope will be published shortly,
Confirms this observation, and shows that all spermatozoa have 2S
chromosomes, while eggs have either 28 or 27.

In B. hirtaria, although there is no ceitain indication as tn which
chromosome is the " sex-chromosome," I think it may be assumed with
some probability that it is one of the larger ones. One of the largest
is always coupled with a small one in the first spermatocytes and such

' J. W. Tutt. Tniii.1. F.iitom. Soc. 1H98, p. 17.

■- R. Gold.schmidt, Zeitschr. f. indiikt. AbsUimm. vii. 1912, p. 1.

2 L. Seiler, Zoul. Am. xli. 1913, p. 24(;.

^ Journ. of Genetics, iii. 1913. p. 1.

J. W. H. Harrison and L. Doncastrr 247

coupling has frequently been described in connexion with hetero-
chromusomes or " sex-chromosomes " ; the size of the chromatin-
nuck'(jlns also indicates that it would give rise to one of the larger
chromosomes. Now all the larger chnjinosomes of hiiiuria are of
greater size than any of zuiutria, so that if the size of the sex-chmmo-
some is any indication of its intensity of action, it is probable that in
the cross zonariu $ x liirturin^ , in which all the spermatozoa have
the large male-determining chromosome of hirtarlu., all the zygotes
might receive a sufficient quantity of the male-determining substance
to cause them to become male, whether they received any from the
female parent or not. In the converse cross, only half the eggs would
receive a male sex-determiner from the hirfaria $, and therefore both
males and females would be produced.

This suggestion does not differ essentially from that used by
Gold.schmidt to explain the results obtained with Lyinantria dispur
and Ij. japniiica, the essence of which is that the "potency" of the sex-
determiner may be different in different species. It is founded, however,
on observations on the size of the chromosomes rather than on the
simple results of breeding experiments. It is admittedly very hypo-
thetical, and does not explain the excess of females which seems
constantly to occur in the cross hirtaria $ x zoiiuria </. I hope to be
able next season not only to amplify my work on the species used this
year, but also to examine other crosses which give comparable results,
and to confirm or disprove the hypothesis suggested by observations on
amplei- and more varied material.

EXPLANATION OF FIGURES.

PLATE XVIII.

All the figures were drawn free-hand, with a Zeiss 3 mm. 1-40 ap. immersion objective
and Compens. oc. 12.

[N.B. In Figs. 19, 22, 23, 26, 27, 30, 31, 32 the reproduction has made some small
chromosomes appear in contact which in the original were near together but quite
distinctly separate.]
Figs. 1 — 14. IS. hirtaria.

Figs. 1, 2. B. hirtaria oogonial equatorial plates. 28 chromosomes in each, if the
small body at the top of Fig. 1 is not a chromosome. There are in each tour
very small and two rather small chromosomes.
Fig. 3. Oocyte nucleus, cut in two section.f. Three chromosomes are cut in both
sections (lettered a, b, c in each). Thirteen elongated chromosomes showing
traces of doubleness, and compound chromatin-nucleolus {ck.n.).
Fig. 4. Spermatogonia! equatorial plate. Four very small and two rather small
chromosomes. Total 28.

17—2

248 Hjihruls of Blittonine Moths

Fig. 5. Early synapsis stage ; fine thread withdrawn to one side of nucleus.

Fig. 6. Later synapsis (bouquet or pachytene stage), with coiled thick thread.

Figs. 7, 8. Primary spermatocyte equatorial plate.s. 13 chromosomes, of which

one consists of a large coupled witli a small (left upper side in Fig. 7, right

lower in Fig. 8).
Fig. 9, «, h. i\ The compound chromosome in side view. In h the small member

appears not to be dividing ; more commonly in metaphase all the large chromo.

somes appear like c
Fig. 10. Primary spermatocyte anaphase ; two daughter groups, cut in successive

sections. In the left group there are 13, one of which is compound ; in the right

group there are 13 with a stained body outside the circle which is probably not a

chromosome.
Figs. 11 — 14. Secondary spermatocyte equatorial plates. In Fig. 11 there are 13, one of
which is clearly compound ; in Fig. 12, none is obviously compouud ; in Fig. 13 the
small member is quite separate from the large ; Fig. 14 an abnormal figure in which
there are 14, one of which is compound.

Figs. 15, 10, 17, 18. B. zonaria.

Fig. 15. Spermatogonial equatorial plate. The chromosomes are so small and

crowded that an exact count is impossible. There are more than 100 and less

than 120.
Fig. 16. Pachytene stage of spermatocyte. The thread is thinner than in liirttirin,

and takes a less typical " bouquet" form.
Fig. 17. Primary spermatocyte equatorial plate. 56 chromosomes quite clearly, of

which four are larger than the rest.
Fig. 18. Secondary spermatocyte equatorial plate. 56 chromosomes, of which tour

are larger.

Figs, ly — 26. Hybrid, zonaria ? x liiiluria ^. (This cross yields only males. )

Fig. 19. Spermatogonial equatorial plate. 14 large chromosomes and about 56

small.
Fig. 20. Early synapsis.

Fig. 21. Thread becoming reticular, apparently omitting the bouquet stage.
Figs. 22, 23. Primary spermatocyte equatorial plates. In Fig. 22, 12 or 13 large

and 50 or 51 email ; in Fig. 23, 13 large and about 40 small ; several of the latter

are of intermediate size, as if consisting of pairs.
Fig. 24. Typical chromosomes from primary spermatocyte prophases, showing

single, equally paired, unequally paired and grouped chromosomes.
Fig. 25. Outline of part of primary spermatocyte spindle in side view, showing some

chromosomes dividing, others apparently not.
Fig. 26. Secondary spermatocyte equatorial plate— about 9 large and 46 small.

Figs. 27—32. Hybrid, hirtaria J x zonaria <? . (This cross gives females and males.)

Fig. 27. Spermatogonial equatorial plate. About 14 large and 55 small chro-
mosomes.

Figs. 28, 29. Early and later synapsis stages, corresponding to Figs. 20 and 21 of
the converse cross.

Figs. 30, 31. Primary spermatocyte equatorial plates. In Fig. 30 about 11 large,
8 or 9 intermediate, and about 40 small ; in Fig. 31, 11 large and 42 intermediate
and small.

Fi". 32. Secondary spermatocyte equatorial plate. About 9 large and 41 small.

JOURNAL OF GENETICS, VOL. IIL NO. 4

PLATE XVIII

'It,—

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••',•

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• • 4

11

• %

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13

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31 "..,«•

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32

ON THE RELATIVE LENGTHS OF THE FIRST
AND SECOND TOES OF THE HUMAN FOOT,
FROM THE POINT OF VIEW OF OCCURRENCE,
ANATOMY AND HEREDITY.

By ONERA a. MERRITT HAWKE8, M.Sc, B.Sc.

CONTENTS.

PAGE

1. The Study of the Foot in the Liviug Subject .... 249

2. The Study of the Foot of the Foetus 257

3. The Relation ot the Toe-types to the Anatomy of the Foot . 258

4. The Inheritance of Toe-types 264

5. Summary of General Conclusions 272

Description of Plates ......... 272

Literature 273

I. The Studi/ of the Foot in the Livinfj Subject.

The following research wa.s undertaken in the hope of discovering
the method of inheritance of two types of feet which were known to
occur commonly in England. As might be expected, however, the work
branched off into a study of the anatomy of the foot, and into an
enquiry into the percentages of these types of feet in the population.

An examination of the living feet of adults and of children, sonn
showed the existence of three types, of which two only are common.
These types are represented diagrammatically in Fig. 1, and photos of
three of the types are shown in Plate XIX.

These types are :

1. (Figs. 1 and 2, i) in which the first or great toe projects beyond
all the others, so that the outline of the toes is a comparatively straight
line. This type will be referred to as L (long great toe). Park
Harrison (1885) stated as a result of examining only 120 persons, that

250

First and Second Toes in Man

5

III

n'

FiR. 1. Diagrammatic Drawings
to sliow Tue-types.

IV

FiR. 2. Toe-tracinns made by metlioil described on page •25.'). I'liotos of
these feet are foand on Plate XIX and Kadiu^raphs on I'lates XX and XXI.

O. A. Merritt Hawkbs 251

the L type was the cuininoiiest and his results are confirmed by my
examination of over 2000 persons. Pfitzner (1901 et seq.) and Weissen-
berg (1895) found this the commonest tjrpe in Germany.

2. (Figs. 1 and 2, ii) in which the second toe protrudes not only
beyond the 3rd, 4th and 5th toes, but also beyond the first. Hence the
outline in this case is an arc. This type of foot will be called 8 (short
great toe). It is much less common than the L type ; but, in spite of
this, it occurs constantly in ancient and modern statues and in nearly
all pictures, although usually in an exaggerated form.

3. (Fig. 1, in) in which the first and second toes are of the same
length but longer than toes 3, 4 and 5. This type is called E. Even
a superficial observer would gather the impression that this type
seldom occurs and careful observation shows it to be very unusual.

The feet of the same individual are usually of the same type, but
not very rarely one foot may be L and the other S. For convenience
the combination of right S and left L is called A (Figs. 1, iv and 2, in),
and left »S' and right L is called B (Fig. 1, v). Combinations of E with
another type have not been met with.

The analysis was made as the result of examining 2-301 persons,
1461 males and 840 females, found in 22 institutes and schools. (See
Table II.)

It was desirable to see the feet of adults as well as of children, but
as adults are curiously unwilling to show their feet, I was obliged very
largely to confine myself to persons under 18. This limitation, however,
need not invalidate the results, for, as will be seen, the proportions are
not materially changed with increasing age.

TABLE J.

(Taken in a simplified form from Weissenberg (18!»5), Table VII, p. 98.)

Ago

5 to 10

11

to 20

1 to 30

31

and over

Number of Persons

198

552

160

00

Right foot:

Percentage

Percentage

Percentage

Percentage

I>II

137

69-2

350

63-4

93

58-1

67

67-7

I<:II

56

28-3

164

29-7

58

.36-3

27

27-3

I=II

0

2 -5

38

6-9

9

5-6

5

5-0

Left foot :

I>II

135

68-2

361

65-4

96

60-0

67

67-7

I<II

58

29-3

1.55

28-1

56

35-0

28

28-3

I = II

5

2 5

36

6-0

8

5-0

4

4-0

Weisscnbi

■rg

(1895) anal

Ivsed,

as shown

in T

able I,

the

feet

of 1009

Jewish children and adults and found that there was no measurable

Firat ami Second Toes in Man

i!7 re to C' -t* -'I X' ro 1-- Oi -^ O '— ' CO re
i-nrirCi— (.xi,— I zr;i— ic^osimcicOi-hO

CO ^-t 1— t 1— I I— I N r-(

[J I -- i I I I I M I 1 I I I I 1 I M I

MM

M M M M M M M "' t

rj M M M ■" ^M I ■» MM M M M
U I -. ^ , I I I I I i -, , I I - - I I - ^.

rt 9

CQ
<

L

M-*-'MM'"l§MM==MMl

OO'MiO I |i-( I I'M lOOi-Hi-H It-tO'M'-Hi-H'-H'^

X O '-' C^ I ^1 CD

I '- M' " I I

^ X r-( ^1 I X OJ

O ^

CO O

OS ^

c-i Tc ■— ' c-i iS

X <:o — c-

a X »-7

C>1

O GO 1 1

X

T*

?r: .-<

o

GO

-% i^ ir:: 'jj I lo -f I I '='1 c-i '^ ^

g ^ -* -1* I I I --I ,x

lO Lf rH O C^ ■— ' ^ CO II

00:1 CI '-T^^"—''^ O^

r-H ^ -M CO •

o t^ -*< t^

rococo-HO^'-<t--»o CO Tj^oi^ic _

^ M I s; M M M M M M M I I I

CO 00 l> > 00 > OS -* GO lo t- (M ^ — X' l>- .X 00 '^ C>1 CO

Ml

a

5

' S5 " „ =

<B i -s J?

X E" -= o -2 = 2

"S *^ ■-■?

5^.S4

r-^" 2 ^ 5 -<

ffi -g

o

a

2 s S

m

„ g 4j d

;5 rf ^ 15 ;i2

n3

"S

h

3

J

m

a

0

ffl

M

<tn

0

O O w

z; s

H 0<

O. A. ^Ierritt Haavkes

253

variation in the comparative length of the toes, on account of age.
But, as the epiphyses unite with the shaft bones of the feet at 17 to
21 years in males and 14 to 18 in females, those ages must be considered
adult as regards the feet, so that the female children examined by me
were approximately adult. During the last four years I have had
19 children under observation from birth. Six of these babies altered
from A, B or S types to L during the first two years of life, but in no
case did a child obviously an L type at birth or slightly later (1 month)
change to any of the other types. Never have I found a change of toe-
type taking place after two years of age. These observations suggest
that the adult condition as regards toe-type is reached by the age of
two years. That a change does take place can be seen by comparing
the radiogi-aph of a child of one (Fig. 3) with the toe-tracing of the
same child at two years (Fig. 2, v). The final position of the bones
depends upon the period at which the child walks, the flexed position of
the bones being retained in the foetal condition until the foot is liattened
by some months of standing and walking.

Fig. 3. Radiograph of child o£ one year, showing S type. (Foot held down by an aJult.)

254 First and Second Toes in Man

The persons examined may be considered as fairly average of the
population, as they came from some schools in Birmingham, two in
Lancaster and a number of institutions in the Midlands, in which the
children were drawn from all parts of England. The analysis shows
the scarcity of all types except L and (S'. It is noticeable that there is
a distinct sex bias in the »S' type, the females being twice as numerous
as the males.

My method of examination was to have the person under observation
stand upright with the mesial surfaces of the feet together. I then
looked down upon the feet, my eyes being as nearly as possible above
the middle of the metatarsal bones. In most cases the toe-type could
be easily and rapidly determined, but, in a number of cases, the second
toe had to be held flat down during the examination and, in some
persons, the first toe had also to be held in its undeformed and primitive
position. My method differed little from that of Weissenberg (189.5),
which he describes thus, " I lay the foot on the ground or on a stool and
hold the leg upright, and after laying flat the toes, observe them from
above in a line parallel with the crista tibia'. This appears to me the
correct method of observation, when the sight falls in the middle of
the foot."

It must be emphasised that what has been studied is the relative
apparent length of the first two toes, oi-, in other words, the protrusion
of one toe be^yond the other. When beginning the research it was hoped
that not only would the relative protrusion be noted, but that exact
measurements of the amount of the protrusion would be obtained.
This was abandoned for several reasons. Measurements on the living
foot could have been made with considerable accuracy by first arranging
the toes in their normal position, but it would have been impossible to
take such measurements, as I was only allowed to examine the feet of
persons in Institutions after undertaking to spend the minimum of time
in the examinati(jn. This compelled me to resort to a personal decision
as to which toe was the longer. In the great majoritj- of cases the
decision is obvious and easy and hence the error from the personal
equation is very small.

Owing to the distortion of the average foot (p. 256) measurements
were quite impossible from the toe-ti'acings (Fig. 2, p. 2.50), upon which
the hereditary part of the work depended. A study of even a few family
trees indicated that it was the actual protrusion and not the amount
of the protrusion which was of fundamental importance and therefore
the lack of measurement was not as important as it appeared.

O. A. Merritt Hawkes -255

As a problem in genetics the question of the inheritance of toe-type
had the interest of being a normal human character, whereas most of
those previously considered from the Mendelian standpoint have been
pathological. I wished to attack the problem simultaneously from
three points of view — first, an anatomical study was made by dissection,
by examination of skeletons, of living feet of child and adult, the foetal
foot and of radiographs : second, statistics were collected as to the
percentages of the various types that occurred, and thii-d, a study of
family histories was made by means of toe-tracings (Fig. 2). In order
to obtain these tracings, 7000 cards were circulated, each card having
printed on it, name (not for publication) and in another corner, right
foot. With the cards went the following instructions:

" I am doing a piece of research upon the human foot, and I should
be very grateful if you wcjuld heljj me by doing as follows : — Place your
bare foot on the enclosed card, and with a pencil held upright draw an
outline of the toes. Do not draw between the toes. One side is tor the
right foot, the reverse for the left. Please pre.ss the second toe out well,
until flat, before drawing, as this toe is frequently not flat owing to
pressure of boots.

I enclose cards also for each member of your family, parents, brothers,
sisters, children and grandchildren.

Please put name on each card and state whether child or adult, male
or female."

There are certain possibilities of error in the drawings: (1) the
pencil may not be held in the same upright position throughout the
drawing; (2) the second toe may not be pressed out flat; (3) the toe
nails may pi-otrude beyond the toes and be of unequal proportional
lengths. The paper did not state the point to be studied, so that there
could not have been any prejudice in the mind of the recorder : particular
directions are given as to how the pencil should be held and there is
small likelihood of the position becoming changed in doing the first two
toes. That the direction " press the second toe out well " was duly
considered, was evidenced by the number of persons who explained that
it was difficult or impossible. In some cases, when the outline was
blurred or uncertain, it has been repeated and then the subject was
requested to note the relative protrusion. As a control <5ver the
accuracy of the drawings, 97 persons (over two years of age) were
examined aftei' their toe-tracings had been made ;iiid amongst these
only one error occurred.

256 First and Second Toes in Man

The paper circularised probably contains one error, viz. the statement
that "the second toe is frequently not flat owing to pressure of boots."
That was the idea held when the work was started, but it is not
supported by observations on the naked feet of children and adults. It
seems probable that the first toe is the only one which is flat in repose,
the flexed position being the normal one for the others, which flatten
out only when in use. The wearing of boots with a stiff sole has much
reduced the use of the toes, so that the tendency is for them to remain
in a flexed position, with possibly a consequent small degeneration of
the toe muscles. These observations bear a certain relation to those of
Pfitzner (1902) on the degeneration of the little or fifth toe. (Plate XXI,
Right foot of Fig. in a).

As a rule, boots are too narrow for our feet, and there is therefore a
lateral pressure which compresses the toes together : but boots are veiy
rarely too short, at any rate among the middle classes, fi-om whose feet
these toe-tracings were principally made. The curled up second toe
cannot therefore be accounted for as a result of foot-gear, especially as
the majority have the first toe longest, and that, because of its very
bulk, determines the length of the boot. The chief deformities are due
to the unnatural curve of the inner margin of the boot, which presses
the great toe outwards and then consequently produces an enlarged
joint. There is a considerable difference in deformity between the feet
of men and women ( Plate XIX, i, ii), the boots of women being by far the
less natural. It is just this lateral displacement that makes the difficulty
in judging the living foot and the tracings. Indeed, the toe-type of
the tracings can only be determined accurately after an examination of
a considerable number of feet, the experience with the living foot acting
as a guide in determining the character of the tracings.

Both <S' and L types of feet occur among races which have never
worn any foot-covering, so that neither the one nor the other type of
toe can be exclusively due to the effects of foot-gear. Flower (1881)
gives some very bad cases of deformed feet but I have not met any foot
whose toes could not be arranged flat except in the case of congenital
hammer toe. The few persons who had this defonned condition were
not enumerated.

In passing, one may note that Park Hari-ison (1883) states that the
third toe is never longer than the second, but this is certainly not the
case as I have found it so in 19 persons. Weissenberg (1895) says
" the third toe longer than the second may be hereditary or due to bad
boots." How such a condition could arise from the latter cause is

O. A. Meruitt Hawkes

257

difficult to conceive, but I have a series nf tracings which seem to
indicate that it is a hereditary character.

Weissenberg (1895) states that the first toe protrudes from 10 to
13 mm., but the second toe seldom more than 5 mm. In these state-
ments I am at one with Weissenberg as regards measurements on 43
persons.

I I'egi'et not having been able to work out the relatiini of toe-type
to height.

11. The Stiuli/ of the Foot of the Foetus.

The only age fiictor of any importance is the foetal period, as shown
by an examination of 91 embryos. (Table III.) This table shows that
the S type is the commoner at this age, as it represents .50 per cent, of
the females and 41 '3 per cent, of the males. It is interesting to observe
that, in spite of the smallness of the numbers, no L type appears in
females until the fifth month, whilst it is found in males of four months.
Perhaps, here too, there is evidence of the female retaining the primitive

TABLE

111.

An

. analysis

of th

\e result

of

' examininy 91 foetuses.

Female Foetuse.s

No.

Malf

: Foetuse.s

No. of Specimens

Age

Type

of Specimens

Age

Type

1

2 months

.S'

2

2

montlis

S

i

3

H

1

3

,,

S

1

4 ,.

S

2

4

,,

S

1

5 ,,

S

2

4

,,

B

1

-' ,,

L

1

4

L

2

6 „

S

1

5

,,

S

1

8 ,,

L

1

5

B

1

8 „

S

0

5

L

3

7—9 mon

tlis

A

2

(5

L

5

7—9

B

3

6

,,

S

10

7—9

S

8

7-

-9 months

S

9

7-9

L

3

7-

-9 ,,

B

3

9 months

H

1

7-

-9 „

A

2

9 „

I.

t)

7-

-9 „

L

1

9 ,.

A

3

9 months

L

1

9

,,

B

1 pail twins

9

,.

A

1

,,

9

,,

S

1 !

set triplets

9

"

B

4-5 =

= 13

i, 23 S,

4 A,

0 B.

46 = 14 L,

19

■5,

3 A, 10

B.

258 Flr^t ((11(1 Si'ctnid Toes in 3Ian

character longer than tlie male. These results by n(( means agree with
the observations of Braiiiie (1*^84), who stated that the second toe was
always the longest in the embryo. Volker (1905) supports this conten-
tion when he writes that, " in all its elements as well as in its pnsition,
the foot of the new-born chilil is analogous with that of the monkey."
P. Harrison (18JS3) examined the iWt of fifteen foetuses (8 male and
7 female) and found them to be 11 L, S E, I S. He does not state
the ages or the relation of the type to the sex. It would be valuable
to have radiographs of foetal feet and babies' feet at intervals from the
sixth foetal month to the age of two, thus showing the change from the
foetal to the adult condition.

The result of the examination of 1+ foetal skeletons is shown in
Table IV, from which it appears that there was not one L type. 'J'his
result is probably incorrect as 11 of the 14 are over six months old.
The error is probably due to the difficulty of preparing and mounting a
foetal skeletal foot, for, to get accurate resiilts as regai'ds the n.'lative
h'ngths of the toes, it would be nfce.s.sary to retain the foot in the flexed
position and certainly not to place it fiat on the ground.

TABLE IV.

Skeletons of Foetuses.

No, of SpeL-imt-ns

Age

Type

1

1

months

S

2

-,

S

3

i;

S

1

7

S

1

7

B

1

8

S

2

9

A

3

i)

S

Total number 14

III. The Relation of the Toe-types to the Anatomy of the Foot.

There has been considerable diversity of opinion as to whether type
L or >S' was the more common. Thus, in the plates of Vesalius' anatomy
(1543) the second toe is generally represented as the longer, but in one
diagram, the two feet (litter. Unfortunately it is impossible to state
how far these diagrams are conventional or actually represent Italian
conditions. Passing on to the " Anthropologia Xova " of Drake in 1707,
one finds the woman of Table XXI and the man of Table XXII both

O. A. ]\Ierritt Hawkes 259

represented with the second toe longest (type .S') but the text reads,
" Pollex pedis is longer than the other toes." In 1864 Vogt, in Lectures
on Man, refers on p. 153 to "the length oi' the great toe which generally
exceeds in man, that of the other toes," but the skeleton on p. 56
shows the second toe as the longest. Marshall in The Human Body
(1875), makes no statement, but the skeleton represented shows tht- S
t}^e and the drawings of the feet are L, S, A and B. Flower in 1881
writes : " The first or inner toe is much larger than either of the others

and its direction is parallel with the axis of the foot It seems to be a

common idea with artists and sculptors as well as anatomists, that the
second toe ought to be longer than the first in a well proportioned
human foot.... Among hundreds of bare and therefore undeformed feet
of children I lately examined in Perthshire, I was not able to find one
in which the second toe was the longest." Braune in 1884 stated that
the second toe was always the longer in the foetus and also among
70 per cent, of adults and this statement is repeated as authoritative by
Stratz in 1903. Kollmann, agi-eeing with Braune, says that 30 per
cent, only of the pojjulation of civilized countries have the L type
of foot, whilst Holden on the other hand says that the majority are of
the L type. Weissenberg in his important paper of 1895 finds that the
majority have the L type of foot. Lazarus in 1896 published complex
tables of the exact measurements of the foot bones, but doi-s not o-ive
their relation to the relative lengths of the toes of the foot. Dunlop in
Anatomical Designs for Art Students (1899) represents the foot and
the skeleton with the second toe decidedly the longer. Pfitzner in his
papers fi-om 1901 to 1903 recognises the existence of both types.
Thomson in Anatomy for Art Students (1906) makes no statement
but writes, " in regard to the length of the toes, there is much diversity
of opinion." Volker (1905) did not examine the li\diig foot nor does he
state the appearance of the skeleton, but makes it clear, that for most
Europeans, the first toe appears the longest. Volker was limited by
examining only the skeleton of the foot, very few of which are prepared
sufiiciently well for careful work. Undoubtedly radiogi'aphs give a
truer idea of the relation of the bones than the dried skeleton. Dwight
(1907) in Variations of the Bones of the Hand and Foot did not touch
on the point now under consideration. An examination, however, shows
both S and L types among his radiographs, but the majority of them
were so taken that no conclusion can be reached as regards the relation
of the toes to one another. From the above summary of the litera-
ture, it becomes obvious that a furthei' enquiry was needed for England.

260 First and Srcoud Toes in Man

The examination of a number of skeletons showed tliat, compared
with the I'esults of a study of the living foot, too large a proportion had
the )S type of fnot.. In <jrder to disco\'er to what this discrepancy is due,
it would be necessary to make an examination of a large number of fi-esh
feet, the fresh skeletons of these feet and the same skeletons when care-
fully dried. As I was only able to carry out this work in one instance,
this question must be left to future investigators.

In consequence of the existence of a numbei' of types of feet, it has
become necessary to discover to what factor the difference in the
apparent length of the toes is due. It was impoi-tant from the Mendelian
point of view to decide whethn tliis difference was due to variations in
length of one or more of the skeletal elements or to the terminal soft
tissue. In other words, is it due to some tmit character or is this
difference, like height, due to a group of characters too complex to
be analysed, or, though due to a complex of factors, may it still be
inherited as a "unit"^

Foi' tile purpose of this I'cscarch radiographs were taken of two
complete families, each consisting of two generations. In one family
there were the two parents and foui- s(jns ; in the other (Fig. 2, and
Plates XIX — XXI), the two parents and three children, two boys and
a gii'l. In both these families the feet of tht' two parents wei-e different,
whilst the children were of various types. The only difficulty whicJi
attended the taking of the photos was in keeping the second toe fiat,
but this was o\'ercome in the case of the young children by means of a
board placed on the toes. The radiographs were taken by placing the
naked foot on a plate and arranging the centre of focus to fall about one-
third fi'om the distal end of the third metatarsal. The arch of the toot
makes the radiogi-aphs slightly incorrect as the distal ends of the meta-
tarsals are nearer to the plate than the proximal ends. As the arch
increases with the length of the bones, that is, from childhood to maturity,
there would be the same pn)portion of error for each foot. Hence the
error may be considered negligible when making comparis(ins between
a series of radiographs, all taken under the same conditions. The only
condition which might have detracted fi-oni the i-esults would ha\e been
a bad case of flat-footedness, but of this, no instance was found.

A comparison of Fig. 2, with Plates XIX, XX and XXI, shows the
correctness of the toe-tracings in these cases. But, it is apparent that the
outline of the foot is not exactly the .same as the outline of the .skeleton.
If the skeleton of the feet as shown by the radiogi'aphs had alone been
examined, the feet would, with one exception (not illustrated), have

O. A. Merritt Hawkes

261

appeared of the same type as indicated by the toe-tracings. In this
exception, the foot when judged by the outline appeared L, but appears
as an E when judged by the skeleton.

Measurements of the thickness of the soft tissue at the ends of the
first and second toes were made and are shown in Table V. These
measurements show that the soft tissue at the end of the great toe is
constantly longer than that of the second toe and that this difference
increases with age. Thus, the average length of the soft tissues for the
first toe is 0'535 cm. and for the second toe is 0'396 cm. On further
analysing the results, it appears that the excess in length of the soft
tissue of the first toe is O'lll cm. for the L type of foot and 0'177 for

TABLE V.

Measurement of Lenr/th uf Soft Tissues at end of First and Second
toes, taken from Badioffraphs.

Tyi)e of Toe

Sex

First Toe

Second Toe

Reference to RafUograpli

>S'

S

0-6

0-.S5

L

,j

OT

0-4

L

5J

0-5

0-4

L

?

0-55

0-3

L

>)

0-5.5

0-4

S

<f

0-5

0-4

S

»)

0-55

0-4

S
L

9
))

0-4
0-4

0-4 1
0-35 1

Plate XXI, III a and iii t

S
S

i)

0-6
0-6

0-4 ^
0-45 1

Plate XX, II a and ii 6

s

t?

0-7

0-4

L

j»

0-6

0-45

L
L

»5

0-52
0-5

0-45 1
0-5 /

Plate XX, irt and li

S

0-4

0-3

Figure 3

L

)>

0-45

0-3

L

J>

0-4

0-35

L
L

0-55
0-55

0-5 1
0-451

Plate XXI, IV a and iv b

L

tJ

0-5

0-45

L

a

0-.5

0-45

S

?

0-65

0-3

S

„

0-6

0-3

L

i

0-5.5

0-4

S

a

0-5

0-45

Average length

... 0-535

0-396

Average leogth

of L

type 0-521

0-41

*

Average length

of S

type 0-554

0-377

18

•2(i:^ First and Second Toes in Man

the »S' type. The importance of these measurements is, that thu L type is
not, at any rate in these cases, due to the extra length of the soft tissue,
and the S type of foot has the second toe longest, in spite of the excess
of S(.)ft tissue at the end of the great toe. These observations by no
means aid in explaining why so large a pni|)(irti(in of skeletal feet are of
the <S' type — except tJiat they suggest that the discrepancy is due to
careless preparation of the feet. Volker met the same difficulty in that
his measurements of the skeletal elements contradicted "les resultats de

jjlusicurs mensurations prises sur les vivants Je dois avouer que dans

ce moment je n'ai pas de donnees pour expliquer cctte contradiction.
Peut-etre faut-il la chercher dans lepaisseur des tissus graisseux au bout
du gros orteil chez les vivants, ou jjeut-etre aussi dans ce fait que
boaucoup de mensurations ont ete faites sans appuyer sur le deuxieme
orteil prcsque toujouis plus plie, pour le redresser en toute sa longueur."
Unless, however, the toes tabulated are unusual, the explanation certainly
does not lie in the soft tissues.

Table VI gives the measurements of the phalangeal bones, of the
metatarsal bones and of the complete colunm or digit, in 26 radiographs.
The measurements have been taken along the median axis of each bone
by means of a transparent glass cm. rule. The axes of the digits are
not naturally parallel to the main axis of the foot. When a digit was
displaced fi-om its correct position, each bone was measured separately
in order to obtain the correct undistorted length.

The most striking result is, that in all cases, except one, the first
phalanges are longer than the second, whether the foot be of the L or S
type. In this one instance, the female of the first family in Table VI,
the bony toes of the left foot are equal and on the right foot the second
toe is 0-1 cm. longer than the first. It is hence obvi(jus that the fiictor
for toe-type does not lie in the toe alone.

Lazarus gives a table with the lengths of the phalanges in 13 cases,
of which 11 show the first toe-bones longer, in one the skeletal toes are
equal, and in the thirteenth case the second phalanges were longer than
the first. He, however, makes no statement concerning the app)earance
of the living foot. Volker, who also dealt only with the skeleton, made
extensive measurements, and he writes : " The first toe in all races is
longer than the second, except it appears among the Melanesians, among
whom the two toes are of equal length."

Table VI shows further, that in every case the second inetatarsal is
longer than the first. Lazarus also found this to be the condition in 1.5
cases, which varied in age from a foetus of 3^ months to an adult.

O. A. Merritt Hawkbs 263

TABLE VI.

Giving lengths in cm. of the Metatarsals (Mt), Phalanges (Ph), arul complete Digit (Dg),
of the Left and Right feet (f ttvo cumjjlete families and of four other persons.

Left Left Left

Toe ^ ' ^ , '■ , . ' N

Person type Mtl ,Mt 2 Mt 3 Mt 4 Jit .'. Phi Ph 2 Ph 3 Ph t Ph 5 Dg 1 Dg 2 Dg 3 Dg4 Dg 5

Parent,? ... B 11 78 7-2 (3 1 ;5-.5 60 oS 51 48 40 13-3 13-9 12-6 ll'l 9-9

Parent? ... .S' 6-.3.5 71 l\-7 (r45 Ir2 49 49 4-1 3-8 32 11-.5 12-1 10-9 104 9-5

Son s 6-8 7-7 7-4 til .5-8 5-7 5-2 4-8 42 3-3 127 13-1 12-6 ll'O 9-2

Son L li-2 7-3 71 t;-7 60 5U 48 4-.5 40 30 116 12-1 117 108 9-2

Son B 6-2 7-4 69 68 51 5-0 47 4-.5 42 3-3 113 123 11-7 111 9-6

Son L 40 4-8 45 44 40 38 3-45 3-3 3-0 26 83 88 84 79 73

Parent j ... L 70 8-4 8-0 7-4 69 6-4 5-8 4-8 4-3 3-7 13-5 143 132 118 ll'O

Parent j ... S — _ _ _ _ .5-3 4-9 4-25 3-7 2-9 _ _ _ — —

Daughter ... A 4-2 4-9 4-8 4o 43 37 3-2 30 27 23 83 8-5 8-2 7-6 7-2

Son L 3-45 3-.55 32 315 3-25 3-3 32 305 25 23 74 75 705 6-6 6-3

Son S 2-5 2-6 24 2-25 22 2-.55 2-4 215 195 185 56 61 55 5-1 5-0

Odd ? ... L 7-8 8-4 82 7-5 72 67 5-8 51 48 3-6 145 144 13-4 12-2 ll'O

Odd;? ... L — _-__— _____ __ _ _ —

Odd s ... L — ____ _____ __ _ _ —

Odd J ... S— ____ _____ __ — _ —

Average all types 5-61 6-36 6-44 5-6 5-13 4-87 4-51 4-05 3-66 3-0 10-72 11-19 10-47 9-6 8-65

Average L types 5-44 6-22 5-97 5-61 5-27 4-84 4-37 3-96 3-55 2-91 10-6 11-03 10-32 948 8-66

Average S types 5-83 6-52 6-12 5-6 4-96 4-91 4-65 4-15 3-77 3-09 10-88 U-38 10-66 9-74 8-64

L type ... 100 114-33 100 90-3 100 1040

S type ... 100 111-88 100 94-7 100 104-6

Right Right Right

Person

Toe
t.vpe

Mtl

Mt 2

Mt3

Mt 4

Mt5

Phi

Ph2

Ph3

Ph4

Pho

Dgl

Dg2

DgS

Dg4

Dg5

Parent,?

B

7-0

7-7

7-1

6-2

5-5

6-1

5-8

5-3

5-0

4-9

18-3

13-75

12-4

11-2

9-8

Parent ?

. S

6-7

7-2

6-9

6-8

6-7

5-0

5-1

4-55

4-0

3-3

11-7

12-4

11-4

10-8

9-!l

Son

s

6-7

7-6

7-0

6-1

5-0

5-5

5-2

4-6

4-2

3-4

12-4

13-0

11-4

10-4

8-6

Son

L

6-8

7-1

6-7

6-0

5-2

5-0

4-8

4.4

8-9

3-2

11-6

12-1

11-1

10-1

8-5

Son

B

6-2

6-8

6-5

5-9

5-0

5-2

4-8

4-4

4-2

3-2

11-4

11-9

11-2

10-4

8-3

Son

L

4-0

4-7

4-3

4-2

4-0

3-7

3-8

3-15

3-0

2-5

8-2

8-7

8-1

7-7

7 1

Parent j

L

7-6

8-3

7-9

7-4

6-6

6-4

5-6

4-6

4-1

8-8

13-9

14-1

12-6

11-6

10-8

Parent ? ...

S

—

—

—

—

—

5-4

5-2

5-0

3-8

3-2

—

—

—

—

—

Daughter ...

A

4-8

4-9

4-7

4-4

4-1

3-65

8-25

2-8

2-32

2-1

8-3

8-9

8-2

7-4

7-2

Son

Son

Odd ?

. L

3-5

8-6

3-35

3-15

8-3

8-5

3-3

8-2

2-7

2-3

7-5

7-7

7-4

6-7

6-56

L

7-95

9-0

8-6

8-0

7-5

6-3

6-2

5-2

4-4

8-8

14-5

15-3

14-1

12-1

11-3

Odd,?

L

6-8

7-8

7-8

7-7

7-3

6-2

5-2

4-8

4-7

3-7

13-1

13-2

13-0

12-8

11-0

Odd ,?

L

6-8

8-1

7-8

7-4

7-1

6-4

5-5

4-6

4-3

4-0

13-3

18-75

12-7

11-8

11-2

Odd ,?

S

6-4

7-1

6-9

6-8

6-6

5-4

5-4

4-7

4-2

3-5

12-15

12-9

12-0

11-25

10-6

Average all types

6-17

6-91

6-58

6-15

5-68

5-26 4-9

4-37

3-91

8-35

11-64

12-18

11-2

10-32

9-25

Average L types

6-24

7-01

6-67

6-21

5-72

5-42 4-94

4-4

4-03

3-48

11-86

12-27

11-4

10-48

9-84

Average S types

6-02

6-7

6-37

6 02

5-6

4-99 4-83

4-33

3-7

3-1

11-13

11-8

10-75

9-96

9-07

L type ... 100 112-3 100 91-14 100 103-45

S type ... 100 111-8 100 96-8 100 106-0

18—2

•264 First and Second Toes in Man

A'^olker, whose work extended to many races besides Europeans, fliinul
that in all races, the second metatarsal was longer than the first.

The next measurements taken were the total lengths of the toe
bones plus the metatarsals, that is, the complete anatomical digit. Here
again my findings agi-ee with Volker, that the second digit (la culumne
metatarso-phalangeale) is in all cases the longer, and this, whetlter the
foot is of the L or of the S type.

The findings from the throe sets of measurements, (1) that the
average length of the first phalanges (bony toe) is always gi-eater than
the second; (2) that the second metatarsal is in all cases longer than
the first, regardless of toe-type ; and (3) that the whole second digit is
constantly longer than the first, throw no light upon why there are L
and S toe-tyjDes. Hence, the relations of the phalanges, metatarsals and
digits have been calculated in percentages, the first digit or any of its
constituent parts being reckoned as 100 (see last two lines of Table VI).
Fnjm these numbers is obtained the imp(jrtant result that both for right
and left feet, the second digit in the S tgpe is proportionately longer than
in the L type. '

IV. The Inheritance of Toe-types.

In (irdt'r to study the inheritance of toe-type, tracings were obtained,
in the manner already described, <if 808 persons (Table VII), of these 381
were offspring of known parents (Table VII, A), 1.50 had one known parent
(Sub-tables B and G), 92 were members of fraternities (Sub-table D),
and 18-5 were the parents of persons in Tables A, B, C, D who do not
therefore appear in these tables. It is at once apparent that the propor-
tion of the various types among these 808 persons is not the same as
among the 2301 persons (Table II) I had personally examined. This
discrepancy, however, is not due to the faultiness of the toe-tracings,
but to the fact that these were made from selected cases, whilst the
feet examined may be regarded as an average of the population. There
were great difficulties in obtaining the tracings, as there are strong
prejudices connected with the naked foot, so that, in order to see the
inheritance in the hylirid generation, very special efforts were made to
obtain families where »S', A and B toe-types were known to exist, and
tithcr families had to be neglected. The large proportion of females
among the tracings is explainable on the same gi'ound, as it was possible
for me to persuade more females than males to submit themselves to
what appeared an ordeal. There is, therefore, for two reasons, selection
and sex, a larger pi-oportion of S type and therefore of A and B types
than would normally occur in the general population.

O. A. Merritt Hawkes

265

TABLE VII.

Table.t A, B, C, D, E, analysimj llic Offspring of Various Miiliuys.
Sub-Table A. In ivhicli botJi parents are Jcnoivn.

Number

of
Matings

Type

of

Mating

, '-

— .

, — ■

— ,

—

— > ,—

-■ — ,

, — ■

— ,

J

?

i

?

<?

? S

?

<?

?

Total Number

54

L i X L'i

85

65

—

—

1

2 1

—

—

—

154

•2

S cT X 'S ?

—

1

2

6

—

— —

—

—

—

9

IC

)L J X .S' ?

18

12

5

3

1 1

2

—

—

43

9

\S i X L ?

10

18

1

—

1 2

1

—

1

37

•25

L^S

28

30

0

3

2 3

3

—

1

-

14

ILs X A 9

13

20

4

1

1 1

1

—

—

42

12

1^1 ^ X L'i

15

20

2

—

2

2

—

—

42

•2(i

L^A

28

40

2

6

1

3 1

3

—

—

—

4

|/, cT X B 9

5

5

—

2

1

— 1

—

—

—

14

1

(Bd X Li

4

3

—

2

—

— 1

—

—

—

10

5

L^B

9

8

—

4

1

2

—

—

—

—

0

\S s X Ai

—

—

—

—

—

— —

—

—

—

—

1

\Ai X Si

2

—

1

—

—

— —

—

—

—

3

1

S^A

2

—

1

-

—

— —

—

—

—

—

2

|S<J X Bi

4

3

-

4

—

1 —

—

—

—

12

3

\Bi X S i

5

1

2

—

—

— 1

—

—

—

9

5

S^B

9

4

2

4

—

1 1

—

—

—

—

0

A S X A i

—

—

—

—

—

— —

—

—

—

—

2

Bi X Bi

1

1

—

—

1

— —

~

—

—

3

1

^As xBi

—

1

—

—

—

— —

—

—

—

1

2

[B i X Ai

2

—

—

—

—

— —

—

—

^

2

3

A^B

2

1

—

—

—

— —

—

—

—

—

0

E S X -E ?

—

—

—

—

—

— —

—

—

—

—

123

—

164

150

11

26

7

8 8

6

—

1

381(190,^ + 191?)

Sub-Table

B

. /

I u

hirh

iiiolltPY Only IS

kiioivn.

Toe Tj

pe of Offspring

L

H

A

B

E

Numbci- Type
of of

. — ■— ,

,—

-■

. '-

-,

.— ■-

-.

.— ■-

— ,

Total
Number of

Mating

s Parent

<? ?

<s

■i

<f

V

S

?

i

¥

Ortspriiig

24

L

16 29

1

7

1

1

—

2

—

—

57

14

S

6 12

1

3

3

2

~

2

—

—

29

5

A

3 5

—

—

—

1

—

—

—

—

9

43

—

25 46

2

10

4

4

—

4

—

—

95 (Sid' +64?)

Number Type

of of

Families Parent

16
1
2

L

S
A

Sub-Table C. In mhich father only is known.
L S A B E

i

19

1

2

?
21

Total

Number of

Offspring

46

1

19 -^

23

55 (25 c? + 30 ?

•2(i(5 Flrxt iiDfl Second Toch In Man

Sub-'T(ible D. Families in which neither mother ur father in knoiru,
designated lii/ H. (homogeneous) and M. (mixed).
L S A B E

of

Number of

Fraternities Type J? <??<?? rf?cf? Ofrspring

21 H. 21 31 — — — — — — — — 52

11 M. 8 13 3 8 2 3 1 2 — — 40

32 — 29 44 3 8 2 3 12 — — 92(35^+57?)

Sub-Table E. Tracings of persons (parents) irho do not appear in

A, Ji, C or D.

L S A n E

' '■ ' ■ ^ Total Xumber

<J? <J? rf? i H if of Offspring

57 57 11 23 13 15 3 (i — — 185 (84 ,f +101 ? )

Sub- Table F. Total of all persons appearing in aboce tables.
L S A B E

' ' ' ■ ' ' ' ' ~ Total Number

t H <J?<f5<??r^? of Ortspring

Total of Group .1 164 150 11 20 7 8 8 G — 1 381 (190,? + 191 S )

i3 25 40 2 10 4 4 — 4 — — 95 (31 J + 04 ? )

C 22 23 2 4 1 3 — — — — 55 (25 <? + 30 ? )

D 29 44 3 8 2 3 1 2 — — 92 (35,? + 57 ? )

E 57 57 11 23 13 15 3 6 — — 185 (84<?+101?)

297 320 29 71 27 33 12 18 — 1 808 (365 <r + 443 ? )

Percentages ^ 8100 7223 8-22 16-02 7-39 7-44 3-28 4-06

Table VII (Sub-table A) gives an analysis of the various types of
matings and of the offspring. The total number of fiimilies consisting
of two complete generations is 123, producing on the average about
three offspring each.

Fifty-four matings of i ^ yi L % were obtained, which pi-oihiced \'i\
offspring, all of which were L except 4, whicli were A .-md B (Fig. 4,
Trees B and C). This indicates that the L type practically bi-eeds true.
The persistence of the L type is seen in Fig. 4 (Trees A, B, C). It
was not possible for me to see the four exceptional cases, but re-tracings
in each case confirmed the type. An enquiry was made into the history
of these four exceptions (3 A and 1 B). Tree C in Fig. 4 shows tlie
family of one male A in which there is no trace of any departure from
the L type. The one case of a male B, as shown in Fig. 4, Tree B,
gives an L ancestry of both the male and female parents, but there is
nothing here to show that one of the parents was not heterozygous.
Consideration may next be given to the two females (Table VII, Sub-

O. A. Meuritt Hawkrs

267

FKiURE 4.

Slmwhiy three family ijroups ( Trees A, B, C) in which there is persistence

of tile L toe-type.

Tree A.

9 = rf

2. 9
L

3.

4.

9 = (?
L I L

9 = 6

L L

9

L

6
L

9

L

9

L

Four generations of L type.
Tree B.

9

L

I

9

L

6

L

9

L

<? = 9

L L

9
L

Two generations of L type with tlio appearance of a B (one of tlie
exceptions mentioned in text).

1.

A

I

9

L

I

i

L

6

L

Tree C.

6 = 9

L

L

1
9 9

L L

i =
I^ 1

- 9 9

1

9

L

I r '

! I

9 i

^ 1^

L

6

L

$ = 9

L I B

-H

I

9
L

6
L

6
L

Three generations with persistence and dominance of i, but the appearance

of an A.

268 First ami Seeovd Toes in Man

table A) of the A type. In one case both parents had L toe-type, but
the tracing of the father was unsatisfactory. Nothing is known of the
ancestry or sibs of the parents. The sister and brother of this case were
undoubtedly L. The second case of a female A is seen in Fig. 5,
Tree D, where an S gi-andparent is known, and where, therefoi-e, the L
father may be a heterozygote.

Unfortunately I only obtained two uiatings of S ^ y. S % ; these
produced nine offsjjring, eight 8 and one L. These numbers are far too
small, but they seem to indicate a tendency to breed true. One of the
families is seen in Fig. 5, Tree E, (iii). This tree shows side by side two
family gi-oups (ii and iii) wliich certainly suggest segi-egation, and here
the offspi-ing are numerous enough to make the results impressive.

The next mating analysed is 8 % x L ^ and L % x 8 ^. These
25 matings gave 80 offspring, divided thus, 58 L,\0 S,\l A and B, 1 E,
or 58 L and 22 of the other types ; here there is partial dominance of
the L over the iS type, approximately 2| to 1, suggesting either that
there are several factors which make up the final result, or that some L
type parents wei'o not pure. If 8 is the ix'cessive type, as appears from
this crossing and others yet to be studied, how can any 8 arise as the
result of the X ^— ^ *S' ci-oss ? The i'Xj)lanation is to be found in the
probability that a few L types are lieterozygous : this would also
account for the few A and B typt's found among the descents of L y. L.
Of the 25 cross-matings, !) only have any iS' offspring, and of these enough
is only known in one case to suggest an explanation. This history is
seen in Fig. 5, Tree F. The grandmother here is an 8, so that the
father may be a heterozygote.

There were 31 matings of Z x ^4 and L x B, producing 124 offspring,
85 Z, 12 ;S', 11 A and B, or 85 L to 28 of the other types, approximately
8|- to 1 ; again the L type shows its dominance, and as would be expected
if A and B are the hetei'ozygous forms, thei'e is a larger projjoi'tion of //
produced than in the L x 8 matings where the L offspring are to the
" other" type offspring as 2| is to 1.

The appearance of any 8 type is not to be accounted for in this
cross, unless again, some L are heterozygous. Amongst the 31 matings
9 only produced the 12 >S' offspring. In 7 of these matings nothing
is known of the ancestry of the L parent ; in the eighth case the L
parent is one of a jjure L fraternity and there is therefore no collateral
history. In the remaining case, the L parent had a pure L ancestry
but had a sister who was A type. It should be noted that of the 12 8
offspring, a large proportion, 10, are females.

O. A. Merritt Hawkes

269.

1.

9
L

9

L

I

i

I

L

9

L

FIGURE 5.
Tree D.

t? = 9

,S' L

I

s

L

I 1

9 9

I i

L L

9

L

I

L

9

I
L

I I

c? = 9 9

B S L

I

S

1.

Three generations «lKi\ving an unexpected A and an expected S in tliird

generation.

(i) I
c? = 9
.1 I L

T

r

-r

Tree E.

~-\ r-

I (ii) I

9 = (?

X I L

I I i I I I

9<:?(?999(?9c?

DLLLLLLLL

I (iii)
? = (?

! I I I I I ! I I
9(?99999(?(?
Z L L S L ,S' .S' ,y ,S'

Two generations to show a hybrid geneiution (i), and pure lireeding in

(ii) and (iii).

Tree F.

1.

2.

3.

i = 9
S

1

9

B

1

c? =
L

9

6
A

S

A

1

i

!

i

9

L

1
1

1

I

6
A

1
1

9

Tliree generations sliowing a \'ariety of types.

.270

First and Second Toes in Man

Only 6 lojitings ti x ^4 and »S' x B were recorded, these produced 24
offspring, 15 L, 7 S, 2 A and B, or 15 Jj type and 9 of the other types.
Again there is dominance of the L type, but the proportion, 15 : 0,
indicates a very real genetic difference between this type of mating and
L X {A or B) in which the offspring are as 3f : 1 or 15 : 4.

The five remaining matings between A and B types gave only (i
offspring, 5 L and 1 A, the L factor being preponderatingly dominant.
The behaviour of A and B with one another and with S and L respec-
tively, makes it clear that they are heterozygous types, just as the
matings L x L and S x S show that these are generally pure types for
the character under consideration.

The net result of all these groupings is to show that the L toe-type
is the dominant, the dominance being, so far as yet determined, irregular.
The usual heteroz5'gous forms are A and B, but some L types also
behave as heterozygotes. That L may bo both a hetero- and a homo-
zygote is seen from Fig. 6, Tree G.

FIGURE 6.

Tn-,' a.

(1) 1

? = c?

/. I n

.1 I /.

! I I /\ I I /\ I I I /\ ! I I !
c?(J?(?99cJ(?99c?9cJ9c?c5??

B L .y L S L L L ,S L L L L L L L L L

Two geuei-atiims to show the two values of L. Amongst the oftspriiig aro
three pairs of orchiiary twins.

Tn-e H.

2.

L

- 9

,S'

\

1

^ ?
A L

1

9 =
s

1 —

! r

= i

A

1

1

1

i

9 =

L

6
L

1

i -

A

= 9

L

1 1

1 1

9 6 -
.s L

= 9

1

n

1

1

1

1

1 1

1 1

1 1

1 1
i S 6
L S L

1 1

S 9

L L

L

L i

? 6

? L

i 1

i $
L L

Three generation.s.

0. A. Merritt Hawkes 271

Consideration must be now given to the possibility that S is some-
times a heterozygous form. The six matings (S' x (A or B) produced
15 X and 9 of the other types, a very large proportion of L, unless S is
sometimes heterozygous or unless the L factor is what may be described
as an excessive dominant, and the matings L x *S' by no means indicate
that. In this connection the family in Fig. 6, Tree H, (i), is interesting,
where the mating >S' $ x J. J" produced two male L. Now this female
S may be a heterozygote, being the child of the mating L ^ y. S $ , or
the two male L of generation three may themseh'es be heterozygotes, a
point which can only be settled during the next decade. This may be
a case in which the heterozygote appears as an S because it is a
female.

There is certainly a sex factor in the S type and this may be
accounted for by the existence of certain female heterozygotes appearing
as 8 type ; on the contrary, there may be a tendency for the male
heterozygote to appear as an L. It is possible that a comparison of the
matings L ^ with A, B or S, and L $ with A, B ov S, may throw some
light on the sex difference. Of the former, there are 34 matings,
producing 72 L, 14 >S', 13 A and B, or 72 L type to 27 of the other types,
a proportion of 16:6. Of the latter (L $ with A, B or ,Sf) there are 22
matings producing 70 L, 9 »S', 10 A and B, or 70 L to 19 other types
or approximately 23 : 6. These results suggest at once that more male
than female L are heterozygotes. Reference must here be made to Fig. 6,
Tree G, in which it is tlie female L which appears heterozygous rather
than the male L.

The above results must be controlled by an enquiry inti > the proportion
of the sexes in the groups of offspring arising from the above two groujjs
of matings. In both cases the A and B types will not be counted as
they show no sex difference.

The mating L J' x (A, B m- S) pi'oduces 86 // and .S' ciffspring,
divided thus: L, 35 </ and 37 ? ; ^^ 3 ^ and 11 ? , i... </ : ? " ^ : 4,
and there are 19'4 per cent, of >S' type.

The mating L $ x (^1, £ or S) produces 79 L and *S' offspring divided
thus: L, 29 J' and 41 ? ; S, 4 cf and 5 ?, i.e. </:?:: 3 : 4, and there
are 12'8 per cent, of 6' type.

The sexes in both cases are approximately the same, but a larger
proportion of S is produced where the L parent is male rather than
female. Thus, the above deduction is confirmed, that when the L type
is heterozygous, it is more frequently male than female.

272 First and Second Toes in Mem

The results of the xaiious niatings may be suiiiinarised thus :

Toe-Types

Type of

/-

A

Mating

LL

ss

A (LS)

B(SL) E

54 LxL

154

—

3

1

2 SxS

1

8

—

— —

25 ix S

58

10

5

6 ]

31 Lx(A

or

C)

85

12

5

6 -

G S X {A

or

B)

15

7

1

1 -

5 AxB

5

—

1

— _

V. Sinnnuiry of Genentl f'ovclnsions.

1. Thivo tiie-types occur, Imt Iwo air iiiiicli iikhi' cininaon ihaii the
other.

2. The Cdmiiionest toe-type in England is one in which tiie first
toe is longer than the others.

3. There is sinne limitation in regai-d to sex, the iS' type occni-ring
moiv counnonly in females than in males.

4. The »S' type of toe is commonest in tiie foetus.

5. The (S' or L types arc pi'iiduccd 1)\' the ccniiparati\e lengths of the
digit, combined with an unknown factor.

6. The L type of foot is irregularly dominant over the >S' t\-pe of

foot.

7. The heterozygotis types are usually A and B, but some L types
are heterozygous.

8. The male heterozygoto tends to be L, the female hcterozygote
to be S.

DESCRIPTION OF PLATES.

PLATE XIX.

Photographs of feet of I. male parent, 11. female parent, III. female child, IV. male
child, outline.^ (toe-tracings) of which are represented in Figure 2.

PLATE XX.

Eadiographs of feet of male (I n and I />) and female (II « and II ')) parents represented
in Plate XIX.

PLATE XXI.

Badiographs of feet of female (Hid and III//) and male (IV k and IV i) children, re-
presented in Plate XIX.

JOURNAL OF GENETICS, VOL. IN. NO. 4

PLATE XIX

JOURNAL OF GENETICS, VOL. ML NO. 4

PLATE XX

Fig. I<(. Bight foot of I. (PI. XIX).

Fig. lb. Left foot of I. (PI. XIX).

Fig. II.(. Left foot of II. (PI. XIX).

Fig. lib. Left foot of II. (PI. XI.\).

JOURNAL OF GENETICS, VOL. III. NO. 4

PLATE XXI

i<l^^ 1I1,(. Kight foot of III. (I'l. XIX).

Fig. Iin-. Left foot of HI. (I'l. XIX).

Fig. IV((. Kight foot of IV. (I'l. XIX).

Fig. IV 6. Left foot of IV. (I'l. XIX).

O. A. 3IBRR1TT Hawkes 27c

LITERATURE.

Anthony, R. (1903 a.) "L'cvolution du pied humain." Dull, et Jlem. Soc.
d'Anthrop. Sur. 5, Tom. in. pp. 818—83.5. Paris, 1903.

(1903 B.) Rev. Scientif. Ser. 4, Tom. xix. pp. 129—139. 1903.

Braune. (1884.) "Ueber einige Formverhaltiii.sse des Jlensehlichen Fusses."

Congres period, inlcniat. des sc. raedicalcs. 8" sess. C'opeuhague.
BuooM, R. (1906.) "On the Ai-raiigemeut of the Epiphyses of the ^Mammalian

IMetacarpals, etc." Anat. Anz. Vol. xxviii. pp. 106 — 108.
Drake. (1707.) "Anatomy" or " Anthropologia Nova."
DuNLOP, James M. (1899.) Anatomical Diayranis for the use of Art Students.

Geo. Bell and Sons, Loudon.
DwiGHT, T. (1907.) Variations of the Bones of the Hand ami Foot. Lippincott

and Co.

Ellis, Havelock. (1904.) Man and Woman. 4tli edition.

Flower, W. H. (1881.) Fashions in Deformitij. Macmillau aud Co., London.

Gruning. (1886.) " Ueber die Liinge d. Finger uud Zehen bei einigen Volker-
stammen." Arch. f. Anthropologic, Bd. xvi. p. .511.

Harrison Park, J. (1882 a.) " On the Length of the Second Toe of the
Human Foot." Brit. Ass. p. 606.

(1883 B.) On the Relative Length of tlic First Three Toes of the Hiunan

Foot." Proc. Anth. Instit. Vol. xill. p. 2.58.

Hasselwander. (1903 a.) " LTntersuchungen liber die Ossifikation des Meusch-
lichen Fusses, etc." Zeitschrift fiir Morphologic und Anthropologic. Bd. v.
pp. 438—508.

(1908 b.) " Ueber die O.ssifikatiou des Fussskeletts." Anat. An:.. 'Qd. wxil.

p. 608.

HoLDEN, " Human Osteology."

Keibel, Franz and Mall, Franklin, P. (1914.) Manual of Hitman Emhrijology.

Klaatsch, Herm. (1901.) " Die wiehtigster Variationen am Skelet der freien

unteren Extremitiiten des Menschen." Wiesbaden, 1901. (Sep. Abdi'. d.

Ergebnisse der Anatomic und EntwiekeUi ngsgcseh . Bd. x. 1900, p. 599.)
Lazarus, J. (1896.) "Zur Morphologic der Fussskelettes." Morph. Jahrbuch,

Bd. XXIV. 1896, p. 1.
Lombroso. (1901.) "Sulla eortezza delP allueo negli epilettioi, nei criniinali e

negli idioti (con 3 figure uel testo)." Arch, cli Psich. p. 337.
Marshall, J. (1875.) The Human Body. 2 vols. Smith, Elder and Co., London.
Oltolesghi e Carrara. (1892.) " H piede prensile negli alienati e nei doliquenti

(con tre figure nei teste)." Arch, di Psichiatria, 1892, Fasc. iv, v, p. 373.

274 First and Secoiul Toiti in Man

PaI'I'ILAl i.T. ■■ L'liuuiuie iiioyuu a Paris." liul/etiH de la Sociele d'Antlirupotogie de

Paris, 1902.
Pfitzner. (1892 A.) "Beitragc zur Keuiitiii.ss des uieuschlichen Extveruitiiteii-

skelets." Sckwalbe's Morphologische Arheiten, Bd. i. 1892.

(1901 13.) "Bcschroibuiig von Aberrationen aus meiiier Samniluug." I). Eid.

Xs.. Ins. Berlin, Bd. xiv. 1901, pi). 113—11.").

• (1903 c.) " Beitrage zur Kenntniss dur Misr>l)ildungen des menschlichcn

E.xtrcmitatenskelets. IX. Eiu Fall vmi bcidcrseitiger Verdopplung dcr
fiuifteu Zebe." Zx. Morph. Stuttgart, Bd. iv. 1902 (pp. 38—393, uiit 1 Taf.).

(1004 D.) " Social-anthroj)ologische Studien. IV. Die Proportionen de.s

erwachsenen Slen.scben." Zs. Morpli. Stuttgart, Bd. v. 1903 (pp. 201—314).

PiER.'iOL, Geo. a. (1907. j Human Ana(om>/. J. B. Lippincott and Co., Phila-
delphia and London.

Str.\tz, C. H. (1902.) La Beaute de la femme, trans, from German by K. Waltz.
Paris, Gaultier, Magnier et Cie.

Thomson, A. (1906.) Anatomy for Art Stiide/its. Clarendon Press, 0.\ford.

Vesalius. Epitome of, 1543.

VoGT. (1864.) Lectures on Man.

VoLKEK, M. Th. (1905.) Variations squelettiques du pied ohez les Primates et
dans les Races humaines. Beaugency, Imprimerie Laffray Fils et Gendre.

\VEissENBER(i, R. (189.5.) " Die Formen der Hand und des Fusses." Zeitschrift
fiir Ethnologie, Berlin, Bd. xxvii. 1895, p. 82.

September, 1913.

THE TEANSMISSION OF SECONDARY SEXUAL
CHARACTERS IN PHEASANTS.

By rose HAIO THUMAS, F.Z.S., F.L.S.

InTR0DUCT1(.)N.

My experiments in pheasant breeding were commenced over eight
years ago after reading Mr Punnett's '• Mendelism " which opened up
a study so fascinating that various schemes were undertaken to
ascertain whether pheasant crosses would follow Mendel's law. Some
of the inter se ' experiments not yet published furnish extremely
interesting evidence of segregation, but every experiment seemed so
involved with the problem of sex (producing unexpected pure recessivea
among small numbers) that for several years schemes have been
arranged breeding back with the female parent or with the male
parent to try and throw some light on the subject.

Cases of the female transmitting the male characters of her species
to her male offspring have been recorded by others previously : amongst
my pheasant breeding experiments many such instances have occurred.

Examples are found in both fertile and sterile hybrids.

Dealing with the first, we will take the Silver $ x Swinhoe ^
series, which form the subject of this paper. In this the F^ male
offspring have to some extent, and F., and Fj male offspring have
the plumage, with the exception of some very interesting mutations
on the under parts^, bulk, call, and moral character (bold and tame)
of the Silver (/ transmitted to them by the Silver hen — the female
parent. In a cross made hetween P. fonuosanus $ x F . versicolor ^ ,
an Fi female, which was in appearance a P. versicolor ^ , yet proved
to have the male Formosan secondary sexual characters present,

' 3id April, 1914. Recent investigations liave shown these to originate from the Male
parent of Fj .

276 Secondari/ Sexual Charactern in Pheasants

triuisinittcd tu her by the female parent, for she transmitted some of
these to her male offspring when mated with a P. versicolor (f .

Turning to the sterile hybrids we find the influence of the female
on the various characters of her male offspring still stronger, a mating
made between P. reevesi $ and P. formosamis ^ produced two F^
males, having the colour, form, pattern and structure of the majority
of the plumage areas together with the bill and bulk of the male of the
female parent species.

A very peculiar case illustrating the phenomenon of the transmission
by the one sex of the secondary sexual characters of the other sex was
sh(j\vn in the difference between the 1910 and the 1911 plumage of
a sterile F^ Reeves x Formosan %. The 1910 plumage vias female,
and much of it was transmitted by the male parent resembling that of
the female of his species. In 1911, the plumage of this -f, Reeves x
Formosan % assumed male characters, and was in every area in which
it occun-ed the male plumage of ihe female parent species, proving that
to this one individual both the male and the female parent had trans-
mitted the secondary sexual characters of the opposite sex of their
species.

A mating between P. reevesi % x P. versicolor ^ produced sterile
offspring, and here also the female parent transmitted to her male
otfsjjring, in several areas, the male characters of her species in bulk,
colour and pattern.

The number of male birds reared in the above crosses has been
considerable, therefore the statement might be made that these facts
are of pretty general distribution in pheasant crosses.

In previous papers {Proc. Zoo. Soc. Ajjril, 1910; Proc. Zoo. Hoc.
September and December, 1912), I have brought to notice facts relating
to the transmission by the male of thc/e/«a^e secondary sexual characters
of his species. In the first, an account was given of a cross between
a Silver ? and Swinhoe ^ , followed by a F^ % y. Swinhoe (^ which
produced an F.. female offspring very difficult to distinguish from a
pure Swinhoe % : and when bred with a Swinhoe </ the only offspring
this F.. female produced was a pure Swinhoe male.

The crossing of Formosan with Versicolor formed the subject of the
other paper, in which it was shown that the male parent transmitted
to his F^ female offspring much of the female plumage of his species
together with the dimension of the i^gg, and that in the F.2 generation
the offspring of F^ Formosan x Versicolor % with P. versicolor J" the
Versicolor male seems to have transmitted every character, bill, leg

R. H. Thomas

277

colour, plumage, habifc and temperament, of the female of his species,
to the F., female offspring, even the dimension of the egg.

It is further worthy of notice that in this experiment, though the
Versicolor male transmitted every character of the female of his species
to his Fo female offspring, yet he did not transmit all his own plumage
to his F„ male offspring.

Following on the above facts, the Gennaeus nycthemerus x G. stuinhoei
series here dealt with confirm the previous investigations into these
strangely sex-limited phenomena, eliminating that certain source of
error, small lUnnbers, and demonstrating that the first were no isolated
cases but evidence of very general application of some significance. In
this exjDeriment, as in all my pheasant crosses, many instances occur of
pattern and colour transference from male parent to female offspring,
or female parent to male offsjDring, or again from one plumage area to
another. Where these have been observed, they have been recorded
to obviate any erroneous classification of these characters as hybrid,
when they are really due, not to a mosaic of the two species crossed,
but to a shifting of colour and pattern fi-om one sex to the other within
one of those species ; but after eliminating these it will still be seen
that in certain of the plumage characters, in the moult habit and in

PEDIGREE OF MATING, 1907.

" 5 " $ Oettnaeus nycthemerus (Silver).
" ^ " (J Gennaetis sidtikoei (Swinhoe).

"iJ" ? X "4" (J
I

Fi ''BA-

Fo

8 birds reared to adult stage

F,

14 birds reared to adult .stage

Fi "BBBA " inter se

10 birds reared to adult stage

I
''BBA'

"5" 9 X "BA" S 1908
I

^ 1

I
"iJ" ? X ''BBA" S 1909

i

I ^ 1

I I

"BBBA" 9 X ''BBBA" S 1911 inter

I I I I I I I I I

Note. Amongst the F^ "BBBA" two mutations, i.e. birds having new characters not
present in either parent species, were bred, a J and a ? which were mated infer se.
Their offspring f 4 inter se ? ? are included in this paper and show clear segregation.
Journ. of Gen. iii 19

278 Secondary Sexual Characters in Pheasants

the leg colour, the F^, F.,, and F^ female offspring are hybrid', notwith-
standing the three doses of Silver $ to one ofSwinhoe J'.

The birds dealt with in this experiment were obtained by the series
of crosses shown in the preceding pedigree of mating.

We will consider the characters of the birds produced by these
matings imder two headings, dealing first with time of maturity,
plumage and leg colour; secondly with the moulting habits and
pattern transference.

1. Time of Matarity, Plumage and Leg Colour.

From 1906 to 1909 it is recorded that the two parent species in
this cross-breeding experiment were, both male and female, immature
at ten months old, and did not breed, but the hybrids F^ " BA " and
F2 " BBA " matured in the first year and bred when mated at ten
months old. There was an increase in size accompanied by premature
sexual development, but not by secondary sexual development. In the
second year as in the jjarent species the adult plumage was assumed.

In 1910, at ten months old, F.j "BBBA" ? $ were penned separately
from F3 " BBBA " (/ cT and they laid no eggs ; but in my pheasant
experiments, extending over seven years, frequent instances have been
recorded of hens not laying when penned apart from cocks in the mating
season, therefore this was not absolute proof of immaturity.

The records of 1911 remark that F^ " BA" 1907 birds mated inte)- se
were laying well, and raising healthy offspring, and that F^ " BBBA "
1909 birds mated inter se were very fertile, hatching 13 birds out of
14 eggs, 12 of which were reared; it is noted that these ^4 "BBBA "
inter se as chicks were strong, healthy, and displayed great courage.

In 1912, during the mating season, the F^ " BBBA " inter se ^ ^
were separated from the F^ "BBBA" inter se (/</ under the same
conditions as F^ in 1910, i.e. kept in adjacent pens with a wire netting
to keep them apart, and though these unmated $ J laid no eggs, being
possibly immature at ten months, yet four of the $ $ attacked and
beat the fifth $ so that it had to be removed, which, occurring during
the mating season, must be attributed to rivalry.

When selecting from the 5 F„ "BBA " (/'(/' in 1909 a male parent
to cross with a Silver $ , the most mixed looking hybrid was chosen,
which in the following summer assumed like the brothers the Silver

' By tlie term "hybrid" is uieaut the blending of characters derived from the two
original parents, due to the simultaneous presence of factors derived from each parent and
patently exhibited in the offspring.

R H. Thomas 279

pattern, but showed Swinhoe colour, in that many feathers were deeply
stained with brown. Amongst the offspring reared from this bird and
the Silver $ (F^ " BBBA," eight in number), two birds exhibited new
plumage characters, " mutations," the one a ^ the other a $ , these two
mutations were mated intei- se in their second year, 1911, and bred the
Fi " BBBA " of this series, reproducing amongst these birds, of which
ten were reared to the adult stage, replicas of the j)lumage mutations.

Two examinations of plumage were made (see Appendices A and B),
the first giving detailed descriptions of the hybrid areas observed
(Plates XXII— XXVI).

In the second examination (Appendix B) I sought, not so much to
define the details of the plumage patterns, but rather to make it my
task to trace their hybrid origin, and in this investigation the immature
eight month plumage of the Silver </, a transition stage between the
chick and adult plumage, proved of great value for it was found to play
a large part in the varied designs of i^„, F^, and F^ in some apparently
hybrid characters, and where absent in other hybrid characters served to
confirm the presence of the Swinhoe characters. It is plain that the
factors for this male immature transition plumage were transmitted by
the female parent to her female offspring, and it is interesting to note
that the characters of the patterns of that plumage show the connection
between the Silver and Swinhoe species : for instance, the flank feather
pattern of the immature Silver (/ (Plate XXIV, fig. 3 b) is very similar
to the breast feather pattern of the Swinhoe $ (Plate XXIV, fig. 2 a, b),
though in colouring they differ widely.

On the wing, tail, breast, and flank of F^, F.,, F.^ and i^j ? ? the
scapular, interscapular and tail covert patterns of the Silver </ eight
month transition plumage can be traced in competition with the breast
and flank of the Swinhoe $ . It is remarkable that upon the breast
and some anterior scapulars of the Silver J transition plumage (Plate
XXII, fig. 16 a, b) was found the same delicate gi'ey shade that dis-
tinguishes the plumage of F^, mutation $ "P" (Plate XXII, fig. 16 c)
and her offspring F^ mutation ? " Cr " a male factor transmitted by the
female parent.

The following list of areas having hybrid characters in F,, F.,, F,,
and Fi may be given : —

Crest (Plate XXII) j Primary and covert

Breast (Plate XXIV) ^"^ [ Secondary and covert (Plate XXV)

Flank „ „ t '1 1 C!entrals

Thigh (Plate XXVI) | Laterals (Plate XXIII)

19—2

280 Secondary Sexual Characters in Pheasants

In the case of leg colour, we also find evidences of hybridism
through all four generations, but in some cases there is distinct
segi-egation of the colour of the parent species. Thus : —

Sil

ver

Parent Species.

[" 5 " $ pale scarlet

(" -B " c/ bright rose

(2 . , \' A" ^ deep dark red

' A" i^ deep dark red

Hyhnds.
(In earlier years, records were not made of every bird.)

Number of birds examined :

(2) Adult F, " BA" ? ? deep bright rose.

(3) Six months old F. "BBA": ? ? (2) bright scarlet, (1) rose
colour.

(26) Chicks F, ''BBBA": (20) like " B" ?, (4) mosaic, (2) like
"A" ?.

(12) Chick stage, five weeks old, F^ " BBBA " inter se : (9) pale
red, (3) mosaic of dull dark red and pale bright red.

(5) Adult stage F, "BBBA " inter se ? ? " GMCKF" bright rose
like the Silver ^ transmitted by female parent.

Thus we see that F^, F^, and F^ generations showed hybrid leg
colour.

The early records in 1911 of Fi " BBBA" inter se notice distinct
evidence of Mendelian segregation even in the chick stage. The leg
colours of the two parent species are as follows: Silver % pale scarlet;
Silver cT bright rose ; Swinhoe % and cf dull dark red. At five weeks
old the leg colour of the 12 F^ " BBBA " infer se is recorded a.s nine (9)
pale bright red legs and three (3) mosaics of dull dark red and pale
bright red, also at two months old notes were made of a strong brown
colour staining the immature plumage of several F4 " BBBA " inter se
? ? and these Swinhoe characters would appear to have been transmitted
through Fj, F„ and F.^, ^ ^ from the one dose of Swinhoe ^ in the first
cross. The hybrid characters of i'^, F.^ and F.^ %% must also have been
derived from the males in each generation.

R. H. Thomas 281

2. Moult habit.

The moult in pheasant crosses is an interesting character to observe.
In two species crossed probably both the order and the season of the
moult will differ. They did so in this series. The Silver moult early
and the central rectrices grow in some time before the laterals. The
Swinhoe moult late and the lateral rectrices grow in completely long
before the centrals appear. Throughout this series all the hybrids
exhibited the mosaic or hybrid moult on body and wings peculiar to
every hybrid pheasant I have reared, quills of young feathers interrupt-
ing the plumage and loose feathers being noticed in the birds long after
the season is past and the moult completed in the two parent species ;
the condition might be attributed to different plumage areas on the
same bird inheriting the late or the early moult character or both, for
new feathers are often shed two or three weeks after arriving at their
full size.

Observations made this year, 1913, on the moult of Gennaeus
nyctliemerus (Silver) (/ and Gennaeus swinhoei (Swinhoe) $ confirm
previous records of the period and order of the moiilt in the two species;
the Silver early, the Swinhoe late, the Silver moult centrals before
laterals, the Swinhoe laterals before centrals. The Silver (f centrals
on July 31st measured two inches longer than any of the laterals, the
Swinhoe ^ centrals measured two inches shorter than any of the
laterals, and were still hidden under the coverts. In the Swinhoe </
the centrals do not grow out till long after the laterals are fully grown.

Ft inter se (F.^ x Fj) Silver x Swinhoe J" " E" has the late Swinhoe
moult and also the Swinhoe tail moult, for all the laterals were shed
by the 21st July, and the centrals were shed on the 2nd August ; these
two moult habit factors must have been transmitted by the Swinhoe
male parent through F^ and F2 males to the F.^ parents. F^ inter se
Silver x Swinhoe $ " G" also shed her laterals first and her centrals
two days later than Fi ^ " E." The dominance of the Swinhoe moult
is also seen in F., inter se of another series, not connected with this
paper.

The phenomena of pattern-transference from one area to another
or from one sex to another, so often seen in pheasant hybrids, occurred
in several individuals of this series, and have been carefully recorded
that these should not confuse the issue by a false suggestion of
hybridism.

282 Secondary Sexual Characters in Pheasants

There is clear evidence that colour and pattern-transference is the
source of the factor for the delicate grey that distinguishes the mutation
F^ "P" ^ and some of her offspring F^, a factor transmitted by the
female parent from the transitional plumage of the male of her species
to her female offspring, transmitted discontinuous and constant to
F3 " P" ^ for the mutation is reproduced in her female offspring
F4 " G" % and also appeared in the immature plumage in three or
four others of her male and female F^, and where it occurred in the
male resulted in a mutation in the breast feathers, which instead of
being like the Silver </ a self black with a dark blue lustre in certain
lights, have down the centre of each feather a long narrow V-shaped
white mark ; the thigh also shows mutation white being present in
considerable quantity.

It is possible that further observation and investigation may reveal
a law governing these phenomena of colour and pattern-transference
and that varieties among species may be found to be partly due to
their influence.

Summary.

Judging from the result of this experiment, it would appear that
the transmission of her female characters by " B" % to her female
offspring was far from complete even in the third generation. In fact
the investigation of the birds concerned j)l<iiiily demonstrates the
following points :

I. Premature sexual development occurred in the crosses, which
bred in the first year instead of like the parents in the second 3'ear.

II. That "B" % failed to transmit her plumage to F, %% " BA,"
and that the hybrid condition oVBA" was continued in /"a ? ? "BBA"
after two doses of " i? " % , and again the same hybrid condition was
observed in i^, $ $ " BBBA " after three doses of " 5 " ? , and finally
when " BBBA " were crossed inter se segregation appeared in the leg
colour and in some plumage areas in i^j ? ^ , and that in others of the
various areas examined the lujhrid nature of the plumage was main-
tained, showing that throughout the series certain of the characters of
the female of " A " parent were carried by F^ , F„ and F.^J'J'.

III. It is also shown that the factors for the male eight month
transition plumage were transmitted by the female parent to her female
offspring and caused a hybrid appearance, but that these sexual hybrid
patterns were quite distinct from the racial hybrid patterns and readily

R. H. Thomas 283

traced to their source : also that the moult habit remained Swinhoe
in the fourth generation, and that the leg colour was sexual hybrid and
racial hybrid throughout the four generations.

IV. That the connection between the two species crossed can be
traced by the eight month transition plumage of the Silver J" and the
adult Swinhoe $ plumage.

V. That a mutation appeared in F^ which has been distinctly
traced to colour and pattern-transference and which was capable of
hereditary transmission (Plate XXII, fig. 16).

APPENDIX A.

Here follow a detailed examination and comparison of the several
plumage areas exhibited in the fi-amed feathers, of which plates are
printed.

Feathers were extracted and set on cardboard from the tail, wing,
breast, flank, thigh and crest, taken from the same locality in each area,
of the following birds. The method emj^loyed was to copy the ring
inscription on to an envelope then extract the feathers and place them
within.

Parent Species.

Gennaeus nycthemerus^, "B" $, Silver J, parent of i^, .
Gennaeus swinhoei, "A" $ , Swinhoe ? , female of parent of F^.
Gennaeus nycthenierus, "Z" $, mottled and vermiculated type of "5."
Gennaeus nycthenierus, " B" <^ , young male of eight months old in iii-st
plumage.

3. F,. "BA" ? ?.

2. F^. "BE A" ? ?.

6. F,. "BBS A" ? ?.

5. F,. {F„ inter se "BBBA") ? ?.

First examination giving pattern and colour in detail.

CREST. (Plate XXII.)
Parent Species.

"B," $ . Crest : blue-black, feathers slender, barbs long, structure
degenerate.

Anterior : length 1^ inches.

Posterior: length 1| inches. Plate XXII, fig. 1.

"B" type of Gennaeus nijcthemerus ? was used throughout the experiment.

284 Secondary. Sexual Characters in Pheasants

"Z," %. Crest : the same as "B" colour, structure, form and length.
"A," $. Crest: brown, ruddy but}' mark down each side of rachis,
feathers broad, barbs very short, structure normal.
Anterior : length f of an inch.

Posterior : length j^ of an inch. Plate XXII, fig. 2.
"B," ^ (8 months). Omitted to extract crest until too late,
adult plumage had grown in.

Hybrids,
"BA."

F,. ? $ . "^" "B," "C," "D," "Er Plate XXII, figs. 3, 4, 5, 6.

Crest of all five hybrid, dark brown, mottled and vermicu-
lated with red brown, feathers broad, structure normal
like female of "A" jmrent. Length of crest the same
in all five.

Anterior : length | of an inch.
Posterior : length 1^ inches.
"BBA."

F,. ??,"4,""5."
Hybrid.

F^. 'i,"A." Crest: same pattern, colour, form, structure and length

as F,. Hybrid. Plate XXII, fig. 7.
F„. ^ , "B." Crest : (rather injured) feathers, darker and shorter.

"BBBA."

F,. ? $ , "A," "B," "P," "C" "^V "B\" Plate XXII, figs. 8, 9, 10.
Hybrid. Crests: similar in pattern, colour and form to F^, the

brown darker, the mottling less distinct.
Structure hybrid between "B" parent and female of "A"

parent.
Lengths of all six similar.

Anterior : length -j^ of an inch.
Posterior : length 1 J inches.

"BBBA," inter se.

F,. ??. (Fainter se),"C," "M:'"F;"'K,""G." AW five hybrid.
Ft. %, "C." Crest: Posterior and Anterior rusty black, mottled ruddy
Hybrid. buff^structure hybrid.

Anterior: length 1 inch.

Posterior : length Ij^ inches. Plate XXII, fig. 13a, b.

R. H. Thomas 285

i^j. $,"J/." Crest: Anterior, brown thickly mottled ruddy buff —
Hybrid. structure same as female of "A" parent.

Posterior black, mottled ruddy buff — structure hybrid.
Plate XXII, fig. 11 0,6.

Anterior : length f of an inch.
Posterior : length 1 1 inches.
Ft. '^,"F." Crest: Anterior, brown mottled ruddy buff, hybrid —
Hybrid. structure hybrid.

Posterior blue-black like "5" parent — structure hybrid.
Plate XXII, fig. 14a, b.

Anterior : length \^ of an inch.
Posterior : length 1^^ inches.
Fi. % , "K." Crest : Posterior and Anterior, brown mottled ruddy buff —
Hybrid. stractuve hybrid. Plate XXII, fig. 12 a, 6.

Anterior : length \i of an inch.
Posterior : length 1^ inches.
Fi. $, "G." Crest: Posterior and Anterior blue-black like "B" parent
Hybrid. but structure hybrid. Plate XXII, fig. 15 a, b.

Anterior: length 1^ inches.
Posterior: length li| inches.
With one exception, pattern, colouration, form, length and structure
of the cre.st are hybrid throughout the "B," "A " series, and that exception,
Fi "0" (Plate XXII, fig. 15), had the structure hybrid.

TAIL. (Plate XXIII.)

Parent Species.

Silver "B," $. Centrals: very finely mottled.

Moults centrals Laterals: first pair vermiculated ; all the others

before laterals. have sloping, irregular bands of alternate black

and white, the black bands the widest. Plate

XXIII, fig. 1.
Silver "Z," ?. Centrals: same as "5" ?.

Laterals: all vermiculated; first and second pairs

a finer, all the others a coarser pattern.
Swinhoe "A," $. Central rectrices : pattern, wide bands thickly
Moults laterals mottled, divided by narrow ruddy buff bands,

before centrals. Plate XXIII, fig. 3 a.

Laterals: copper brow'n, inner margins mottled

black, tips black. Plate XXIII, fig. 3 b.

286 Secondary Sexual Characters in Pheasants

Silver "B," </ (eight months bird). Centrals: coarse vermiculation
ground white tinted fulvous.
Laterals : sloping regular bands of black and white of
equal width. Plate XXIII, fig. 2.
F^. "BA," ??. Tail. Laterals: pattern of female of "A"
parent centrals. Pattern-transference. Plate XXIII.
fig. 4.

F.-.. "BBA," $?. Tail. Centrals and laterals : %6?-mZ. Plate

XXIII, fig. .5.
F^. "BBBA," ^^. Tail. Centrals: mutation.

Laterals : hybrid. Plate XXIII, figs. 6 and 7.

" BBBA" inter se.

Ff. $, " C." (F3, inter se.) Centrals: pattern same as "B" $
first pair of laterals, colour hybrid. A Pattern-
transference.
Laterals : liybrid pattern and colour.

Fi- %, "M." Centrals: pattern of ".^" ? centrals; colour, hybrid.

Laterals : hybrid pattern and colour.
F^. $ , " F." Centrals : hybrid pattern and colour.

Laterals : 1st and 2nd paii-s unpattemed olive brown —
hybrid. Other laterals same liybrid pattern and
colour as centrals.
Fi. %/'K." Centrals: /;?/6?-icZ pattern and colour.

Laterals : somewhat resembling " B" $ and young

male " B " in pattern ; colour, hybrid.

Fi. ? , " G." Centrals : (mutation) unpatterned \ Like parent

olive gi-ey. I F, ? " P,"

Laterals : Very finely vermiculated [ " grey muta-

grey. tion."

Of these J^4 ? ? (F, "BBBA," inter se), " M," " F," "K" and " G"
are darker and gi-eyer bii'ds than any in this series and could be picked
out easily when laid amongst the " B" ? parent, the i^i ? ? , the i^j ? ?
and the i^s ? ? not even excepting the grey mutation parent ? " P."
The heads and necks are dark grey, but inter se " BBBA " F^ $ " C "
is a somewhat lighter bii-d than her four sisters, being much the same
colour as her parent "gi'ey mutation" "P" only a lighter and more
olive tint of gi"ey.

R. H. Thomas 287

BREAST AND FLANK. (Plate XXIV.)
Parent Species.

"B," $. Breast: fine, irregular, broken, zigzag lines of white on
a black ground. Plate XXIV, fig-, la, b.
Flank : pattern the same, lines fewer and coarser, ground
black. Plate XXIV, fig. 1 c.
"Z," $. Breast: very fine mottling on brownish olive gi'ound.

Flank : mottling coarser, ground lighter tint of same
colour.
"A," $. Breast: coarse black lines of painted form on ruddy buff
gi-ound. Plate XXIV, fig. 2«,, b.
Flank : same ground colour, lines irregular and broken.
Plate XXIV, fig. 2 c.
"B," cT, transition plumage. Breast and Flank. Plate XXIV,
fig. 3 a, b, c.

Hybrids.
"BA."

F,. %%, "A," "B," "C," "D" "E," all hybrid in colour and jjattern.

Plate XXIV, figs. 4, 5.
Fi. $$, " E" and "B." Breast and Flank coarsely vermiculated,

somewhat resembling "B" parent but finer lines.
Fi. $ $, "G" and "A." Breast and Flank, black vermiculation on

ruddy buff ground.
"BBA."

F,. $ $ , "^ " and "B." Hybrid colour and pattern.

^"2. $, "A." Breast and Flank very fine grey vermiculation on pale

buft' ground. Plate XXIV, fig. 6.
F2. $ , "B." Breast and Flank, coarse black vermiculation on pale buff

ground. Plate XXIV, fig. 7.

"BBBA."

F3. %%, "A" "B," "C" and "P" are again all hybrid in pattern and
colour and show a remarkable degree of variation in
these areas.

F^. %, "A." Breast and Flank alike, coarse, broken, black, zigzag lines
on a light buff ground.

288 Secondary Sexual Characters in Pheasants

F3. $, "B." Breast: coarse vermiculation, ground a pale olive shade of
buff; the Anterior feathers resemble "Z" $.

Flank: coarse vermiculation or broken lines on same
ground.
F.,. $, "C." Breast: Posterior feathers, in that the lines are light on
a black gi-ound, resemble "B" parent, but in the form
and colour of them, pale buff, they differ. Anterior
feathers hybrid. Plate XXIV, fig. 8a, h.

Flank: light lines on black groimd resembling " B" parent
but again the form and coloiu', jiale buff, differ. Plate
XXIV, fig. 8 c.
F,. ?,"P." (Grey mutation.)

Breast : Anterior dark grey ; Posterior, fine dark grey
vermiculation on pale grey ground. Plate XXIV,
fig. 9 a, b.

Flank : same as Posterior feathers.

"BBBA," inter se.

F,. %%, "K," "F," "M" "G," "G," all hybrid, patterns widely diver-
gent, showing segregation.
F^. ^,"K." Breast and Flank: light broken lines on dark ground

like "B" parent are nevertheless hybritl in pattern

and colour of lines, which are buff. Plate XXIV,

fig. 12 a, 6.
Fi. $, "F." Breast almost an unpatterned olive of a buff shade, some

very fine mottling on part of the feathers. Plate XXIV,

fig. 13 a.
Flank fine vermiculation on lighter shade of buff' olive.

Plate XXIV, fig. 136.
Fi. $,"il/." Breast: Anterior hybrid; Posterior like "A" ? also
Pattern- resembling in pattern the flank feather of the young

transference. eight months old male of " B " parent. Plate XXIV,

fig. 36, c. A sex pattern-transference. Plate XXIV,

fig. 14 a.
Flank : hybrid ; over all these feathers lies the buff stain

oi"A" parentage. Plate XXIV, fig. 146.
Fi. ?, "C." Breast and Flank: hybrid, like "M," the "A" parentage

is clearly traced both in pattern and in the colour, a

warm buff, which suffuses both areas. Plate XXIV,

fig. 10 a, 6.

R. H. Thomas 289

F,. % , "G." (This bird is being bred from, 1913.)

Breast : Anterior, dark silver grey ; Posterior, dark silver
grey vermiculation and mottling on lighter grey ground.
Plate XXIV, fig. 11a.
Flank: Dark silver grey mottling on slightly paler grey
ground. Plate XXIV, fig. 116.

Degeneration in size of feathers.

It is to be remarked that the size so diminishes and the form so
alters in the breast and flank feathers of two F^ and all the Fi as hardly
to be recognised as having been extracted from the same areas as those
of the females of the parent species. The pointed form referred to is
a male character in both species but much more extreme in the
Swinhoe cT.

Laying the four i^^ M(<e»- se ? $ "A"," "F," "M,""G" breast uppermost
beside "B" parent and the female of "A" parent, the Mendelian segre-
gation is distinct, $ "K" resembles "B" parent, $ "F" neither parent,
whilst $ $ "M" and "C" though hybrid, yet have a strong fixmily
resemblance to the female of " A " parent.

WING. (Plate XXV.)

Parent Species.

Primaries. Secondaries. Major Coverts.

"B," $. Primary: inner vane grey self, outer vermiculated buff
Primary Major Covert : similar.
Secondary : grey ground vermiculated and mottled buff.

Plate XXV, fig. 1.
Secondary Major Covert: similar. Plate XXV, fig. 10a.
"Z," $. Primary: same as "B," colour browner, vermiculation
finer.
Major Covert : to correspond.
Secondary : same as Primary.
Major Covert : rather browner vermiculation.
"A," $. Primary: wide bands of dark grey divided by narrow
bands of warm ruddy buff
Secondary: similar pattern, bands of warm ruddy buff
narrower.

290 Secondary Sexual Characters in Pheasants

In both areas the ruddy buff bands are spotted here
and there with dark grey. Plate XXV, fig. 3a, h.

Majar Coverts : same colour and pattern with heavier
sjjrinkling of spots on the ruddy buff bands. Plate
XXV, fig! 106.
"B" (/■ (eight months old). Transition plumage.

Primary : grey ground, coarse fulvous vermiculation on
outer vane and tip of inner vane.

Secondary : grey ground very coarse fulvous vermiculation
on both vanes. Plate XXV, fig. 2 a.

On both the lines of vermiculation are generally con-
tinuous and unbroken.

Adult Secondary. Plate XXV, fig. 2b.

Hybrids.
"BA."

F,. ? ?, "A," "B," "C" (Plate XXV, fig. 4), "D," "E," all five have

both Frimaries and Secondaries the pattern and colour

of the female of "A " parent : the ruddy buff bands are

rather more thickly spotted.
Major Coverts, both primary and secondary gi-oups the

same as the female of "A" parent. Plate XXV, fig.

10 c, d, e.
"BBA."
F,. ? ? , "A," "5" (Plate XXV, fig. 5). In both birds Primaries and

Secondaries colour of female of "A" parent, pattern

hybrid, a coarse broken vermiculation, both vanes

patterned.
Major Coverts of both hybrid.
"BBBA."

F,. ? ? , "A," "B," "C," "P" (grey mutation), also "A'" and "B\"
F3. $ ?, "A," "B," "J.'," "B^." Primaries and Secondaries, colour of

female of "A" parent, pattern hybrid, similar to F.^.

Plate XXV, fig. 6.
F^. $,"P" (gi'ey mutation). Primaries and Secondaries, very dark

grey finely mottled. Plate XXV, fig. 7.
F^. %,"C." Primaries and Secondaries. Colour of female of "A"

parent but with finer vermiculation than "A," "B,"

"A\" "BK"
In all, vermiculation is much broken.

R. H. Thomas 291

"BBBA" inter se.

F,. "C""M,""F""K:"'G."

F^. %,"K." Primaries, Seccmdaries, Major Coverts, hybrid, same as

F^ "A," "B," "C," "A\" "B\"
Fi. $ , " F." Primaries, Secondaries, Major Coverts, hybrid, greyer

colouration more finely vermiculated. Plate XXV,

fig- 9-
F^. %, "C." Primaries, Secondaries, Major Coverts similar to "K" and

F,. Hybrid. Plate XXV, fig. 8.
Fi. ^,"M." Primaries, Secondaries, Major Coverts, pattern similar to

Ft "C" and "K," but less distinct and colour greyer

and darker. Hybrid.
Fi. $ , "G." Primaries, Secondaries, very dark gi'ey finely mottled like

i^3 $ " P " the parent.

THIGH. (Plate XXVI.)

In this area in the pheasant the dimensions of the feathers are
reversed, the Posterior being the smaller, the Anterior the larger.

Parent Species.

"B," 5. Thigh: Posterior and Anterior, pure white zigzag lines

on black gi'ound. Plate XXVI, fig. la, b.
"Z," $. Thigh: Posterior and Anterior, pattern grey, less regular

sometimes undefined on cream ground.
"A," $. Thigh: Posterior and Anterior, dark grey zigzag lines on

ruddy butf ground. Plate XXVI, fig. 2 a, b.
"B," cT (eight months old). Transition plumage. White lines on

greyish black ground. Plate XXVI, fig. 3a, b.

Hybrids.
"BA."

F,. %%,"A,""C,""D,""E."

Thigh: patterns hybrid, some like "Z," colour ruddy bufi

same as female of "A" parent. Plate XXVI, figs.

5, 6 a, b.
F^. %, "B." Thigh: the lines are finer, more delicate and so broken

as to be almost mottled. Plate XXVI, fig. 4 a, b.

292 Secondarfi Sexual Characters in Pheasants

"BBA."

F... %%,"A,""B." Thigh: colouration of i^i, all feathers smaller than

parents "B" and "A" or F^.
F.,. %,"A." Thujh: pattern %6W(i, somewhat coarse. Plate XXVI,

fig. la, h.
F._. $,"5." Thigh: pattern .same as F, " B," hyhrid. Plate XXVI,

fig. 8 a, h.

"BBBA."

F,. ? $, "^," "B," "C" "P." Thigh: all feathers very much smaller

than parents and F^. Patterns and colouration hybrid,

much individual variation.
F3. $ , "C." Thigh : Posterior and Anterior closely resemble "B" parent

in pattern, ground hybrid' tinged buff. Plate XXVI,

fig. 11 (T, b.
F-i. ?,"P" (grey mutation). Tliigli. : Posterior and Anterior fine grey

lines on paler grey ground. Plate XXVI, fig. Xla, b.
F-i. %,''A." Thigh: Pi)sterior same as i^;, $ "P," Anterior same as

F^ "G," but ground a deeper tinge of buif'. Hybrid.

Plate XXVI, fig. 9((, b.
F3. %, "B." Thigh: Posterior and Anterior pattern and colour same

as F„ "A." Plate XXVI, fig. 10a, b.

"BBBA," inter se.

F,. ??. "C," "M," "F," "K," G." In all these birds the thigh
feathers are still more diminished in size than those

OfP3??.

Ff $,"('." Thigh: Fosterior, pattei-n of female of " A" parent, ruddy
Degeneration buff ground paler. Anterior, pattern of female of ".4"
in size of parent, ruddy buff ground paler. This feather measures

feather. just half the length of that of female of "A" parent

extracted from the same zone.
Fi. $,"ilf." Thigh : Posterior and Anterior, pattern of female of "A"
Pattern- parent breast, ground colour pale ruddy bufil Pattern-

transference, transference.
Pj. % , "F." Thigh : Posterior and Anterior, grey pattern like "Z" but

ground colour hybrid of the ruddy buti" shade belonging

to "A" parent.
F^. %,"K." Thigh: Posterior, pattern of " B" parent. Anterior,

feathers very degenerate but "B" pattern can be

traced, ground hybrid, pale fulvous.

R. H. Thomas 293

Fi. $,"(?." TJiigli: Posterior and Anterior, pattern of very marked

scratchiiigs of dark silver grey on pale cream gi'ound.

Size of feathers very smalk

Tile gradual degeneration in tlie dimension of the feathers in the

thigh area (Phxte XXVI) is a marked and distinct feature of the series

which reaches the point in Ft "K," of degeneration of the barbs. In

this area of their plumage the F^ $ ^ (F^, inter se) show the same

individual Mendelian segregation as in the breast and flank (Plate

XXIV). This dwindling in size associated with change of form was also

observed in the breast and flank of some -fa $ ? and all the -f 4 ? ?

(Plate XXIV).

APPENDIX B.

Investigatiux into Origix of Hybrid Coxditiox.

TAIL. ( Plate XXIII.)

Central putter n Lateral pattern

F,. "BA," ^^Jlybrkl. Silver (/, 8 month.?, Swinhoe ?, central.

lateral. Plate XXIII, fig. 2.

Culoiir Colour

Swinhoe $ , central. Silver $ , lateral.
vSilver ? , lateral. Swinhoe $ , lateral.

F,, F„ F„ all hybrid. Plate XXIII, figs. 5, 6, 7.

^3 $ mutation "P." 2nd lateral exact shade of delicate grey found
on Silver J' transition jjlumage anterior scapulars. Plate XXIII, fig. 7
and Plate XXII, fig. 6.

BREAST AND FLANK. (Plate XXIV.)

Second examination alongside of 8 month immature plumage of
Silver J'.

Patterns

F,. " BBB A" inter se,"C" %, Hybrid. «. Silver $, "£." Plate XXIV,

fig. Ic.
b. Silver J', 8 month. Plate
XXIV, fig. Sb.
Breast. Plate XXIV, fig. 10((.

Flank. „ „ b.

Colour

Fig. 10 «, b. Swinhoe $ and
„ Silver </, 8 month,

Journ. of Gen. in 30

■2i)i Secondary Sexual Characters in PJieasants

BroHst feathers. The prnnted form of Silver j" adult breast and
Swinhoe </ adult breast feathers. Size degenerate.

Patterns
F,. "BBBA " inter se, "M" ?, Hybrid. Swinhoe ?, PL XXIV, fig. 2.
Plate XXIV, fig. 14, a, b. Silver ^ , 8 month, PL XXIV,

fig. :i
Silver ?, "/>" variety, PL ■
XXIV, fig. 1.

Colour

Silver c/.
Swinhoe $.

Breast feathens : the pointed form of Silver (/ and Swinhoe J'

breast feathers. Size tlegenerate.

Patterns

Fi. " BBBA" inter se, "K" $ , Hybrid. Silver j", intcrscapsulars tran-
sition.
Swinhoe $ .
Silver ?,"£."

Culour
Silver (/, transition.
Swinhoe %.
Breast feathers : the pointed form of Silver (/ and Swinhoe </
breast feathers. Size degenerate.

WING. (Plate XXV.)

Fi. "BA,' % %. All very .similai- and level in colour, pattern, and form

Hybrid. of Primaries, Secondaries and their respective coverts.

Both Primaries and Secondaries are alike in pattern.
Numbei' of bands : same as Silver (/ adult secondary (Plate XXV,

fig. 2b). Pattern, Swinhoe % (Plate XXV, fig. U). C'o/ow.r, Swinhoe ?.

F.,. "BBA," $ $. Pattern, colour and form of Primaries and Secondaries
alike in all the birds. Resemblance closest to Silver cf
8 month transition secondary (Plate XXV, fig. 2«) with
a possible trace of Silver $ secondary (Plate XXV, fig. 1)
and Swinhoe ? lateral pattern (Plate XXIII, fig. 36).

F-j. "BBBA." No change, hybrid and very level in colour, pattern,
and form; replicas of F„ "BBA" ? ?. Plate XXV,
figs. 6, 7.

R. H. Thomas

295

Finding that two distinct patterns of the Silver hen exist in the
aviaries of this country I sent a breast feather of each to the Natural
History Museum, South Kensington, and I append to this paper a table,
kindly made at my re(iuest by Mr Ogilvie Grant, of the patterns of the
wild G. Kijctheineniv from the specimens at the Natural History Museum,
which shows these birds to be in a mutating or extremely varying

Notes froji British Museum, Natural History,
Ornithological Department.

Gennaeus )iijcthemerus.

? Ah Cb'uug, Fobkien, S. China
? Kuatum, N.W. Fohkien
?

. ?
?
S
?
?
?
?
? Anioy (Aviary)

B, Z, see illustiatiou Ijelow (1) B type, (2) Z type.

C, Lateral tail -feathers finely mottled or vermiculated with black and white.
i>. Lateral tail-feathers black and white in wavy bars.

B

z

c

D

7 Dec.

'02

B

—

c

—

26 Nov.

'01

B

—

—

D

16 Mar.

'99

—

Z

—

D

3 Nov.

'98

B

—

c

—

8 Nov.

'98

—

z

—

D

6 Nov.

'98

—

z

—

D

8 Nov.

'98

—

z

—

D

6 Nov.

•01

—

z

c

—

3 Nov.

'98

—

z

c

—

17 Oct.

'96

-—

z

c

—

Jan.

'98

—

z

—

D

0.1^

Fig. 1.

Fig. 2.

Breast feathers of the two types B (1) and Z (2) of the female Gennaeus nycthunerus.

20—2

296 Seconrlart/ Sexual Characters in Pheasants

Cdiiditiiin (if plumage, of such a nature that the two extremes of the scale
of patterns would undciubtedly. if found in the cTcTi have been classed
separately. Now though these might account in some measure for the
widely divergent patterns met with in the series we are considering, yet
they do not furnish us with the real clue to the origin of those colours
and patterns found in the legs and plumage oi' "BA," "BBA," "BBBA,"
and inter ae "BBBA" $ $ ; thus, in one area, the crest (which has the
same structure, pattern and colour in both "B" and "Z" varieties of
Silver $ ), the characters of the female of "A" parent can be clearly
traced in every adult ? throughout these four generations, and in
several other plumage areas (with the exception of one or two of the
birds) the characters of the female of " A " parent are present.

"Wild females of the Silver Pheasant vary considerably individually
in the characters of the outer tail-feathers and the feathers oi the breast
and flanks.

As regards the outer tail-feathers, in some specimens they are
black with oblique mottled lines of white ; in others they are very
finely vermiculated with black and brownish-white ; and all intermediate
gradations are to be found.

Young birds have the breast and flank-feathers uniform brown ; in
older birds the breast and flank-feathers may be either finely or coarsely
marked with black and white.

These remarks are based entirely im wild shut ImitIs from China."

DESCRIPTION OF PLATES.

PLATE XXII.
CKEST.
FiK. 1. "B" Silver ?.

Fig. 2. "A" Swinhoe ?, t'Binale of ".J" parent species.
Fig. 3. ''lU" f\, "A" {.
Fig. 4. " BA" 1<\, "U" ?.
Fig. 5. "«.l"i''i, "D" 9.
Fig. 0. "BA" t\, "E" ?.
Fig. 7. "BBA" F.,, "A" ? .
Fig. 8. "BBBA'- F:t, "A" 9 .
Fig. 9. "BBBA" Fa, "C" ?.
Fig. 10. "BBBA" F:,, " P" ?.
Fig. 11. "BBBA " inter se. Ft ,
Fig. 12. "BBBA" inter sc, F^,
Fig. \S. " BBBA " inter se. Ft,
Fig. 14. "BBBA" inter se, F4,
Fig. 1.3. "BBBA " inter se, Ft,

Fig. IG. "B" Silver ^ (male of "Jl" pareut species) transition plumage. 11, Breast;
li, Scapular.
" BBBA" Fs. "!"• ?.

Mi'

? . Anterior, n ; posterior, h.

!{•'

' f . Anterior, 11 ; posterior, h.

C"

9 . Anterior, a ; posterior, /).

p"

9 . Anterior, a ; posterior, h.

'G''

'(mutation). Anterior, a; posterior, h.

"I

>"' pareut species) transition plumage.

c,

Lateral lectrix.

R. H. Thomas

297

PLATE XXIIl.

TAIL.

Fig. 1. "iJ" Silver ? (" B " parent). Lateral.

Fig. 2. " B " Silver s (transition plumage). Lateral.

Fig. 3. " A " Swinhoe ? , female o£ " A " parent species.

rectrix.

Fig. 4. " BA" Fi, " B" i . Lateral.

Fig. 5. " BBA " F2, "A " ? . Lateral.

Fig. C. "BBBA" F3, "P" f . Lateral.

Fig. 7. " BBBA " F-,, ".I " ? . Lateral.

o, Central rectrix; h, lateral

PLATE XXIV.

BREAST AND FLANK.

Fig.

1. "

Fig.

2. "

parent

Fig.

3. "

Fig.

i. "

Fig.

5. "

Fig.

6. "

Fig.

7. "

Fig.

8. "

Fig.

9. "

Fig.

10. "

Fig.

11. "

Fig.

12. "

Fig.

13. "

Fig.

14. "

i? " Silver ?. Breast: n, posterior; t, anterior, i-, Flank.

J "' Swinhoe ?. Breast: o, posterior; 6, anterior, c, Flank (female of "A'

species).

B" Silver (J (transition plumage), a, Breast, i, f, Flank.

BA " Fi, " B" ? . Breast : a, anterior; h, posterior.

BA " Fi, "E " ? . Breast, posterior.

BBA " F-2 , " A " i . Breast, posterior.

BBA " F2,"B" ? . Breast, anterior.

BBBA"F3,"C" ?. Breast: «, posterior; ?<, anterior, c, Flank.

(I, posterior ; h, anterior.

Breast, n. Flank, h.

Breast; a, posterior; fc. anterior.

Breast, a. Flank, /*.
Breast, a. Flank, h.

Breast, a. Flank, h.

BBBA " Fi,"P" 1 . Breast :
BBBA " inter se, Fi, "C" ¥ .
BBBA" inter se, F^, " G" ? .
BBBA " infer se, Fi, "K" S .
BBBA" inter se. Ft, "F" ? .
BBBA" inter se, Fj, "31" ? .

PLATE XXV.
WING. SECONDARIES AND MAJOR COVERTS.

Fig.

1.

Fig.

2.

Fig.

3.

Fig.

4.

Fig.

5.

Fig.

G.

Fig.

7.

Fig.

8.

Fig.

9.

' B " Silver ? . Secondary.

' B " Silver i . a. Transition secondary : '), Adult secondary.

' A" Swinhoe S , female of ''.1'' parent species, a. Secondary; h. Secondary.

'BA"Fi, "C" ?. Secondary.

Fig. 10.

"BBA" F.,, "B" S.
"BBBA" F:5, "B" !
" BBBA" F3, "P" ?
"BBBA" inter se, F^,
"BBBA" inter se, F^,
Major Coverts. a,"B

Secondary.
Secondary.
Secondary.
" C " ? . Secondary.
" F" ? . Secondary.
' Silver } ; h, " A " Swinhoe ? ;

c,d,

'BA" Fi,^ f .

298 Secondanj Sexual Characters in Pheasants

PLATE XXVI.

THIGH.

"i"' Silver ? . a, posterior; 6, anterior.

"J " Swinhoe ? , female of "J " parent species. «, posterior; h, anterior.

"iJ" Silver i (transition plumage), a, posterior; /), anterior.

"BA" F^, '• B" ?. n, posterior; 6, anterior.

"BA" t'l, "C" ? . a, posterior; h, anterior.

"BA" Fi, "£" s . a, posterior; h, anterior.

"BBA" Fri, "A" ? . a, posterior; }i, anterior.

"BBA" F^i, " B" f . a, posterior; ii, anterior.

"BBBA" J"':!, ''A " ? . a, posterior; b, anterior.

"BBBA'' F-i, " B" ?. a, posterior; h, anterior.

"BBBA" F-j,, "C" ? . a, posterior; fc, anterior.

"BBBA" Fj, " P" ?. a, posterior; /<, anterior.

Fig.

1.

Fig.

2.

Fig.

3.

Fig.

4.

Fig.

5.

Fig.

0.

Fig.

7.

Fig.

8.

Fig.

9.

Fig.

10.

Fig.

11.

Fig.

12.

JOURNAL OF GENETICS. VOL. IlL NO. 4.

16

PLATE XXI

1

West.Ne-wmaii chr.

JOURNAL OF GENETICS, VOL. IIL NO. 4

PLATE XXIII

JOURNAL OF GENETICS, VOL. IIL NO. 4

PLATE XXIV

^^:i^V^^

JOURNAL OF GENETICS, VOL. IN. NO. 4

PLATE XXV

JOURNAL OF GENETICS, VOL. IN. NO. 4

PLATE XXVI

<o

-N

Sitw

^

J^
^

,#'

oo

r-

i

HEREDITY OF MELANISM IN LEPIDOPTERA.
Bv W. BOWATER, B.D.S.

In certain of the Lcpidoptera (not naturally black) black specimens
occasionally appear, and it has been noticed that this melanism has
become more and mcjre frequent of recent years, that in some species
the black specimens are to-day distinctly more numerous than the
original type, and in some localities have even superseded it'.

Such an evolutionary change obviously possesses features of con-
siderable scientific interest, and much has been written on the subject
of its causation and the exact method in which it is being carried on,
but the question is still unsettled.

Apparently the term melanism as applied to Lepidoptera should be
restricted to the substitution or increase of black on the wings or body,
or both, at the expense of some other colour ; but any darkening of the
ground colour, even if not black, has been included in a general way,
although strictly the term melanochroism should be applied to the
latter.

I have compiled a list- of 211 species of British Lepidoptera in
which melanic or melanochroic specimens have been recorded.

Although the heredity of melanism in the higher orders has been
worked out in many species, comparatively little seems to have been
done with regard to Lepidoptera, and it is still a fairly open question as

1 Collective Inquiry as to Progressive Melanism in Lepidoptera. L. Doueaster. Fyiit.
Record, Vol. xviii. 1906.

2 (a) Melanism and Melanochroism. Tutt.

(b) List of varieties and aberrations peculiar to British Lsles. Tutt. Ent. Record,

1902.

(c) Melanism in Yorkshire Lepidoptera. Porritt. Trans. Brit. Ass. 190G.

(d) Feuille des Jeunes Naturalistes. April, 1913.

{e) Standfuss, Handbuch d. Gross-Schmetteriinye, 1896.
if) Les Uleciinismes du Melanisme et de VAlbinisme cliez Us Lepidopteres. Pictet.

Geneva, 1912.
(u) Entoinulogist and other sources.

30(» Hereditji of Melanism in Lepidoptera

to whether the heredity of mehmism in these insects follows definite
rules, and especially as to whether it follows Mendelian Laws of
Heredity.

In the literature bearing on the subject records are to be found (if
the breeding by entomologists of many thousands of moths which have
melanic forms, but the accounts are very seldom of such a nature as to
allow inferences to be drawn as to whether the heredity is Mendelian
or otherwise.

In 12 species definite investigations have been made or are being
made :

1. Spilosonid lubricipeda. The e.^periraent of Mr J. Harrison' on
this species and its melanic variety zatima, Cramer (= var. radiata,
Haworth), and other experiments made here-, and abroad', seem to
show that there is no complete dominance, but by selective breeding-
pure strains of the melanic forms have been obtained, and an interesting
point is that these forms become larger and stronger than the type, and
double-brooded.

2. Aplecta nebiilosa. Experiments have been carried out by
Messrs Mansbridge, Harrison and Main (see table), and to my mind
prove that melanism is a Mendelian dominant; and a point of special
interest is that the homozygous and heterozygous forms are probably
distinguishable. In the first instances it seemed certain that the
melanic variety thompsoni, which is black but has white terminal area
(i.e. wing margin and fringes), is the homozygous form ; and the melanic
variety rohsoni, a dark suffused form, at times almost black, but never
with white terminal area, is the heterozygote.

Mr Mansbridge now considers it likely that only those specimens of
rohsoni which have grey scayiular areas are the heterozygotes and
hopes to settle the question by experiments now in hand.

Of the 10 families recorded in the table, only one, the last, fiiils to
support the assumption that melanism is dominant, the fact that
13 type specimens appeared seems to show that both parents are
heterozygous, which is not impossible, as entomologists differ as to the
exact differentiation of the two melanic varieties, thompsoni and
rohsoni; and unless the specimens are in perfect condition it is most
ditticult to decide.

1 Eutomoloiii.tt, 181)3, pp. i;5, 247, 340 ; 1894, pp. 9-5, 129, 20.5.

"- Xaturalist, 1894. W. Hewett, p. 253.

'■' Standfuss, HiouU/iich il. iiross-Schmetterliniif, 1890, p. 307.

W. BOWATER

]01

Another difficulty is that this species is particularly liable to
succumb in confinement even in the hands of the most expert, and
inbreeding seems always fatal.

Ap/ecta nfljiilosa.

Observer

Parentage

Melanic

Date

Tltompsoni

Robsoni

Type

<? X ?

1908

Harrison '

Type X Type

—

—

all

1905

Mansbridge

Type X Type

—

—

76

1906

„

? X rohsoiii wild

—

16

15

1908

,,

Type X thompsoni

—

10

—

1909

,,

Type X rubsoni

—

4

1

1909

robsoni x robsoni

7

15

8

1910

.,

wbso7ii X robsoni

15

27

9

1911

robsoni x robsoni

3

27

25

1908

Har:

rison and Main '

rolisoni x robsoni

*67

158

75

1910

2

thompsoni x robsoni

16

21

13

* Approximate numbers.

The melanic form does not appear to be increasing in Delauiere
Forest, and of 972 imagines reared by Mr Mansbridge from larvae col-
lected there during the last 9 years, 142 are melanic, but only 3 of
these are thompsoni.

Thus it seems that although melanism is dominant in this species,
yet in Nature there is some factor which counterbalances this at present,
and prevents the variety becoming more common.

3. Triphaena comes. Messrs Prout and Bacot^ conducted breeding
experiments in 1903, 1904, which showed that in this species melanism
is dominant, but not completely so, as its depth varies, but Mendelian
segregation occurs.

4. With Boarmia repandata, Mr Mansbridge has been experimenting
for some years, but ijuite conclusive results are not expected until
next year.

5. The same applies to Tephrosia consonaria and Mr J. W. H.
Harrison.

1 Proc. Ent. Soc. Land. October 21, 1908, p. Ixi. Proc. South Lond. Entom. mid Nut.
Hist. Soc. 1908—09, p. 84, and 1909—10, p. 04.

- Trans. Ent. Soc. Lond. May 3, 1911.

3 Ent. Record, Vol. xv. 1903, pp. 217—221 ; Vol. xvi. 1904, pp. 1—5. Proc. Ent. Soc.
Lond. 1905, pp. 67—71.

302 Heredity of Mela n lam hi Lejtidopfera

6. Acidalia virgularia. The classical experiments of Messi's Front
and Bacot' on this species, where they reared 10 generations represented
by 5531 specimens, showed that the two varieties — London, dark, and
French, light — were not Mendeliaii f'oniis of the species, but the experi-
ments were of great value in proving other points in heredity.

Ova fi-om pairings in generation F^^ were given to Mr Bateson,
who in February, 1909, handed the resultant lai-vae to Mr W. B. Alex-
ander-, who successfully reared them and carried on further broods, and
eventually, with the aid of the former specimens and his own, demon-
stratt'(l that although A. virgidaria and its light variety, cunteneraria,
had been proved not to be Mendelian forms of the species, yet one of
the differences between the two forms, namely, the sjDCckling in rinjn-
laria, behaves as a Mendelian dominant to the absence of speckling in
canteneraria. Here then is a Mendelian rule, but the variation in the
ground colour obscured the evidence of it.

Mr R. T. Baumann* has recently bred black specimens of virgidaria
and with Mr Bacot is conducting further experiments on the heredity
uf this form.

7. Acidalia contiguariu. Mr W. Buckley, in a series of exjDeri-
inents on the dark form of this species extending through 8 generations,
has produced evidence, conclusive to my mind, that the dark form is
a simple Mendelian dominant. The experiment is still going on, and
detailed results are as yet unpublished.

S. Xanthorhoe ferrugata = Goremia unidentaria. The breeding
experiments of Mr L. B. Prout^ in 1895 show conclusively that this
species follows Mendelian rules.

In addition to the facts contained in the paper, the author Lew
certain conclusions which he afterwards modified, and is now quitj in
accord with the statement that the experiment proves the reddish ( r
purple form to be dominant and the black form recessive''.

9. Abraxas grossuluriata. The black variety of this moth {varle-
ijata) has been bred by Messrs Porritt and Newman, and it seems that
it follows Mendelian lines but is recessive to the type".

1 Proc. Roy. Soc. Loml. B. Vol. 81, 190'J.

2 Proc. Roy. Soc. Land. 1912, p. 4.5.

3 Entomologist, February 1913, p. 30.

' Tran>: City of Loud. Knt. Soc. 1897—98, pp. 26—34, and J'cuh.--. Eiit. Soc. Land.
1906, pp. 525—531.

= Proc. Ent. Soc. L(vid. 1907, p. xx.
'■ Eiit. 1909, p. 75.

W. BOWATBR 303

Mr B. H. Crabtree is now conducting an experiment, commenced in
1912, which apparently makes this certain.

10. HemeropMla abruptaria. Mr Harris' and Mr T. H. Hamling-
have shown that the dark form is a Mendelian dominant.

11. I can discover only two records of experiments on Odontopera
bidentata which throw light on the heredity of its melanic form : —

a. A paper by the late Mr T. H. Hamling^ in which it is recorded
that a black $ taken at Methley deposited ova, which produced
70 black and 66 type imagines. From this family inbred, 14 successful
pairings are reported :

-L

Parentage

Imagines

Male Female

Melanic

Type

i broods combined

T X

T

24

44

3 broods combined

T X

M

2.5

13

3 broods combined

M X

T

37

10

4 broods combined

J/ X

M

90

6

Total

249

specimens

These results can be explained, but only in a way which would be un-
justifiable were it not that a much more extensive experiment has now
been made, which supports the following argument :

The first family was mixed, therefore neither parent was dominant.
The original $ parent being black must therefore have been hetero-
zygous; and the </ either similar; or type (rece.ssive). In the former
case, the offspring should be 75 per cent, melanic (heterozygous and
homozygous) and 25 per cent, type ; and in the latter, 50 per cent,
melanic (all heterozygous) and 50 per cent. type.

The comparative frequency of occurrence of the two forms in nature
makes it more likely that the J' was type ; and the fact that 136 moths
were reared from the 146 ova deposited, makes this almost certain,
because the melanic form seems to be the more hardy, and of the
10 ova which foiled to reach maturity, the majority would probably
be type, and even if all were black, the numbers would only be 80 : 66.

Thus it seems that all the 70 blacks were heterozygous, so we should
expect M X M to give 75 per cent. M and 25 per cent. T, but the
mortality amongst the larvae was very high indeed, and would account
for the hardier melanic outnumbering their type brethren by more
than 3:1.

» Proc. Ent. Soc. Lonil. 1904, p. Ixxii.

2 Trans. City of Land. lint. Soc. 1905, p. 5.

'■> Trans. City of Land. Ent. and Nat. Hist. Soc. 1903, pp. 40—43.

304 Heredttji of MeJcuiism in Lcpidoptera

In the results of the type x melanie crosses tlie same explanation is
probable.

With regard to the inelanic specimens recorded from parents both
type, it seems that one of the 4 J 's had really paired with a black (/.
Pairing occasionally occurs immediately the wings have dried, and all
breeders of Lepidoptera know how difficult it is to exclude all possi-
bilities of error when dealing with large numbers of pupae. On the
much less probable of the t\vo original hypotheses, viz., that the
original (/ parent was melanie (hetei-ozygous), some of the 70 would be
homozygous, and this would the more easily explain the preponderance
of melanics in their offspring.

h. A paper entitled "Melanism in Yorkshire Lepidoptera," by
Mr (i. T. Porritt'.

A melanie female captured in 1904 deposited ova, which developed
into 6 melanie and 3 type sjDecimens. From the black moths a large
brood was reared, which included 75 per cent, black specimens, and from
these again a considei'able number were reared in which the percentage
of black was even greatei-.

Mr Porritt has also repeatedly ci'ossed tyjje and melanie and found
the pi'oduce is about half and half of the two forms. These results are
easily reconcilable with the following experiment and the conclusions
drawn therefrcmi :

In the autumn of 1009, on the advice of Mr Leonard Doncaster,
I connnenced an experiment on Odontopera bidentata in the hope of
discovering whether the heredity of the melanie form of this species
followed any rule or law.

I commenced with a perfectly open mind on the subject, and in fact
had only a slight acquaintance with the various facts ,-uid theories of
heredity and variation.

In November, 1909, 1 obtained 12 pupae, from which in April, 1911,
6 type and 6 melanie moths emerged. (Family 09.1).

Two pupae were also obtained from a different source and produced
one type and one melanie .specimen — 09.2 (see 1909 table).

Six pairings were made, but in only two cases were the offspring
successfully carried to maturity: — 10.2 and 10.3.

In June, 1910, I (jbtained 24 larvae bred from two melanie parents,
probably both of 09.1 flimily. The resultant moths ai'e 10.4.

' Trtiiis. Brit. Ass. lyOC.

W. BOWATER 305

Oclontopera bidentata. 1909 T'able.
Imagines

Parentage
Male X Female

Label of
family

Type

jMelanic

Totals

Male

Female

Totals

Male Female Totals

M X il

09.1

4

2

6

4

2 6

12

M X il

09.2

—

1

1

—

1 1

2

Total

14

0.

bideittata.

1910 Table.

Label of
family

Imagines

Parentage
Male X Female

Type

Melanic

Male

Female

Totals

Male

Female Totals Totals

09.1 r X 09.1 T

10.2

1-5

23

38

—

—

—

38

09.1 T X 09.2 T

10.3

22

21

43

—

—

—

43

il X i[

10.4

—

—

—

10

7

17

17

il X il

10. -5

—

1

1

2

—

2

3

Warwick-
shire 1

York-
shire 1

2

—

—

—

2

Total for year 103

0. bidentata. 1911 TaMe.

Imagines

Type Melanic

Label of

Formula Parentage family Male Female Totals Male Female Totals Totals

\10AM X 10AM 11.11 _ _ _ 12 11 23 23

■ "1 10.4 ill X 10.4 Ji 11.1.5 „ _ _ 16 10 26 26

Totals

28

21

49

49

c.

) 10.3 T X 10.4 i)/

11.4

22

16

38

38

1 10.4 M X 10.3 r

11.13

—

—

—

28

31

59

59

Totals

50

47

97

97

rlO.S .1/ X 10.2 T
\ 10.2 T X 10.4 il
U0.4 31 X 10.2 T

11..5

9

14

23

11

11

22

45

E.

11.0

12

22

34

21

17

38

72

11.8

—

—

—

—

2

2

2

Total

Is ...

21

36

57

32

30

62

119

1^10.2 r X 10.2 T

11.1

5

4

9

—

—

—

9

F.

J 10.3 I X 10.2 T

11.12

3

2

5

—

—

—

3

j 10.3 T X York. T

11.14

1-5

14

29

—

—

—

29

I War. r X 10.3 T

11.23

20

26

46

—

—

—

46

Totals ...

43

46

89

—

—

~

89

Yorkshire

—

1

1

2

1

2

3

5

Middlesbrough ...

11.24

22

1-5

37

—

—

—

37

Birmingham

11.25

4

1

5

—

—

—

5

Cambridge

—

2

1

3

—

—

—

8

Gateshead

—

—

4

4

—

—

—

4

Bm'nley

—

1

1

2

—

—

—

2

Total

56

Total for

year

410

306

Heredity of Melanism in Lepido2)tera

0. bidentata.

1912 1

able.

Imagines

Type

Melanic

Forniu

la Parentage

Label of
family

Male

f'einale

Totals

Male

Female

Totali-

Totals

A.

11.0 -U X York. .1/

12.44

—

—

—

7

5

12

12

or
B.

Wakef. 31 x Wakef. M 12..52

—

—

—

6

3

9

0

Totals

13

8

21

21

■ 11.1.5 .1/ X 11.0 7'
11.11 .1/ X 11.23 T

12.23

—

—

—

21

19

40

40

c.

12.34

—

—

—

31

35

66

66

11..5 T X 11.15 il/

12.41

—

—

—

8

6

14

14

Totals

60

60

120

120

rll.4 M X 11.15 il/

12.11

3

3

6

11

14

25

31

11.5 M X 11.0 M

12.16

7

0

16

25

17

42

58

11.13 M X 11.13 M

12.33

2 + G'+2

4

8

8

16

20

11.13 .1/ X 11.13 .1/

12.40

4

6

10

16

14

30

40

D. •

11.0 il X 11.9 J/

12.46

2

1

3

7

3

10

13

11.5 M X 11.5 il

12.18

—

—

—

1

1

2

2

11.11 .1/ X 11.0 M

12.29

1

—

1

1

1

2

3

11.13 il X 11.9 .1/

12.53

—

1

1

6

8

14

15

ai.13.1/ X 11.13.1/

12.57

—

—

—

—

'2

2

2

Totals ...

19

22

41

75

68

143

184

11.4 il X 11.24 T

12.12

11

7

IH

15

18

33

51

11.24 T X 11.15 il/

12.20

5

0

14

13

9

22

36

E. ^

11.13 .1/ X 11.14 T

12.26

21

25

46

17

17

34

80

11 15il/ X 11.14 T

12.26 C

9

4

13

4

1

5

18

11.11 il X 11.14 /'

12.31

8

8

16

8

3

11

27

11.9 il X 11.9 r

13.39

11

4

15

13

5

18

33

Totals ...

65

57

122

70

53

123

245

Burn. T x Burn. T
11.24 r X 11.5 T

12.7

1

—

1

—

—

1

12.10

20

20

49

—

—

—

49

11.24 r X 11.5 T

12.17

36

25

61

—

—

—

61

11.24 r X 11.24 T

12.24

4

10

14

—

—

—

14

11.25 r X Gates. /'

12.43

22

21

43

—

—

—

43

11.9 T X 11.9 T

12.45

21

ii;

37

—

_

—

37

F. '

11.23 r X 11.24 T

12.47

7

2

0

—

—

—

9

Camb. r X Camb. T

12.50

15

7

22

—

—

—

22

Tunb. /' X Tunb. T

12.50

10

7

17

—

—

—

17

I. w. r X I. W. T

12.63

46

61

107

—

—

—

107

Mosel. 7'x Mosel. T

12.64

32

30

62

—

—

—

62

Wimbl. /• X Wimbl.

T 12.65

5

10

15

—

—

—

15

*■ Burn, r X Burn. T

12.66

8

6

14

—

—

—

14

Totals ...

227

224

451

—

—

—

451

Total for year

1021

Total of specimens shown

1548

* Gynandromorph.

W. BOWATER

307

In September, 3 j^iipae from an altogether fresh .source were ob-
tained (10.5).

The following April they emerged, 103 specimens (see 1910 table).
Various pairings and cross-pairings were made, and 13 families safely
reached maturity — 410 .specimens (see 1911 table).

From the.se 62 pairings were made, and 33 families, comprising
1021 specimens, were raised to maturity (see 1912 table).

From the specimens which emerged in 1913 various pairings were
made, including crosses between the palest of the type forms and the
most nearly black, and at the present moment there are over 1000
living })upae, comprised in 21 families, 4 of them represented by over
100 specimens each '.

0. bidentata. Classified Tahh.

Specimens

Families

Type

Totals

(A)

12.44 or 12.52 or both

2

—

21

21

(B)

10.4 10.5

2

—

20

11.11 11.15

2

—

40

12.44 or 12.52 or both

2

—

21

Totals

6

-

90

90

(^)

11.4 11.13

2

—

97

12.23 12.34 12.41

3

—

120

Totals ... 5

(D) 12.11 12.1(3 12..S3 12.40

12.46 5

12.18 12.29 12.53 12.57 4

217

39 = 24.1°/„ 123 = 75.9 °/„

2 20

217

Totals

(E) 11.5 11.9 11.8 3

12.12 12.20 12. 2i; 12.26f

12.31 12-39 6

41 = 22.3% 143 = 77.7 7o 184

57 62

122

123

(F)

10. 2

11.1

12.7

12.43

12.-59

10.3
11.12
12. 10
12.45
12.63

Totals

11.14
12. 17
12.47
12.64

9

11.23
12.24
12.50
12.65

12.66

13

179 = 49.17% 185 = 50.83% 364
81

451

Totals

621

621

Combined Totals

48

841

635

1476

' 470 imayines have emerged up to date (March 14, 1914) and they support the former
evidence.

308 Hereditji of Melanism In Lepidojttera

I have given all care to the technique of the experiment such as
profuse note taking (including many points besides those referred to in
this paper), frequent counting of the specimens in all stages, double
labelling, and so on ; and Mr Doncaster has kindly made \arious visits
of inspection during its progress, and is willing to accept it as reliable.

In a survey of the specimens reared in the exjjerinient, certain
points are worth noting :

(1) 111 melanic specimens the aliddiiieii, legs, and antennae are black
or nearly so; and the wings are usually black, but in some cases have
a slight brownish suifusion as in (12.52): and in sdine cases a patch of
tan on the fore-wing (as in 12.11).

Type specimens have light coloured alxloiiien, legs, and antennae.
The wing colour varies between light brownish-yellow and dark brown,
and many northern .specimens tend to lie banded, but all show the
normal markings, and are always easily distinguishable at a glance
from melanic specimens.

(2) In both type and melanic forms the male specimens are darker
than the corresponding females ; there is no other sign of influence of sex.

(8) I cannot jjoint out any difference to the eye between homozygous
and heterozygous forms.

Now if we classify the results from the jioiiit of view of the constitu-
tion of the parents we note that :

(1) Type by type breeds typi; only, no iiu'lanic specimen cropping up
among the 621 type specimens bred (F).

(2) Segregation is clearly shown in !) fairiilics(l)) where 2.5 per cent.
of type specimens appear in the otfsjjring of melanic parents.

(3) Five families (C) are cases of a pure homozygous melanic paired
with a type, giving all heterozygous melanic offspring.

(4) Nine families (E) are cases of a heterozygous melanic pairing with
a type, the resultant progeny consisting of type and melanic specimens
in equal numbers.

(5) The fact that the melanic specimens are usually slightly in
excess of the expected percentage seems to show that they are more
hardy and few'cr die during the early stages, and this is bi>rne out by
the fact that the melanic .specimens usually emerge more early in the
year than their tyjDe brethren, and will even emerge in temperatures
between 1° and 5° C.

W. BOWATER

309

(6) This species stands close breeding well and inbreeding fairly
well, though in many of the families which died out the cause seemed
to be inbreeding.

Genealogical Table. 0. bidentata.

1907

Wild M T

"V

1908

Mmy. Mm

1909

MM+Mm+T

MM+Mm+T

11 n 11.15 11 4 11.13 U 9 11.1 11.12

MM -t Aim rnUMm MmMmAlmtT T T

1912

11.11

'"----/ Middleaboro

1123 115 11.24

12.34 1231

Mill. Mm +T

12.20

Mm+T

15 46 12.26C ■. .,,, mC „ 12 12 „

T Mm + r ^^+ ^" + '^ Mm i /• ","

12.33 12-39 f'2 2« '"'"

MM+ Mm\Mm + T Mm 4- T
+ T j

Thus each of the 50 families (48 bred from ova) falls under one or
other of the following well-known Meudelian formulae :
If MM = homozygous melanic,

M^n = heterozygous melanic,
mm = type,
then MM x MWI = MM (A)

MM X i¥m = AIM + Mm (B)

MMxmm = Mm (C)

Mm X Mm = MM + Mm + mM + mm (D)
Mm X mm = Mm + mm (E)

and mm x mm = mm (F)

I feel that the foregoing is sufficient evidence to prove, that in
Odontopera bidentata, melanism follows Mendelian rules and is dominant.
Journ. of Gen. iii il

310 Heredity of Mehmism in Leindopieni

VI. The question of the exact method of the heredity of melanism
in the fomiliar Peppered Moth, Amphidcisys betularia, is of special
interest because the melanic variety, doubled ay ( tria . in the 65 years
since it first appeared has multiplied and spread all ovei- England, and
is now far commoner than the type, and is often ijuoted as a good
example of a distinct alteration in a whole species occurring in our own
times and apparently not due, directly at any rate, to man's influence.

In the Type form the body and wings are white, abvmdantly dusted
with black. On the costal margin are five black spots. From the first
two arises a duplicate angulated black first line, to the third is attached
a black discal sjjot, the fourth <jriginates the second line, which is angu-
lated. On the hind margin is a series of black dots, extending into the
white cilia. Hind wings white (especially so in costal area), dusted or
spotted with black, and having usually 2 or 3 okscure partial transverse
lines. Central spot black. Leg tufts brownish grey, tarsi and antennae
in female black spotted with white. In male antennae brown-black.

In the Melanic form (doubledayana) the whole insect is almost
uniformly smoky black, except lower portion of face which is white, and
a round white spot at the extreme base of the costa of each fore-wing.

Varieties of the type have occurred where the definite markings
have been reduced, leaving the whole white surface but .scantily dusted
with black, and one specimen has occurred almost white. In other cases
the typical markings are accentuated and somewhat increased.

In what are usually termed the intermediate forms, the normal
markings are much increased ami become partly confluent, scj that in
many cases the moth is black spotted all over with white dots and
minute irregular blotches. The hind-wings are almost always lighter
than the fore-wings.

Between the type and the donbledayaria variety every gradation of
intermediate has occurred, although tlie intermediate is the rarest of the
three forms. It occurs in nature as well as in bred specimens. The
thorax and abdomen are of exactly the same colouration as the wings in
all specimens of type and varieties.

The black which is velvety in a live specimen diminishes in intensity
on death, and has a strong tendency to " rust " on the slightest provoca-
tion, and after a time even in spite of precautions.

The accompanying table consists of those records of genetic value of
breeding this moth which I have been able to find.

In February, 1910, I started an experiment which I hoped would
furnish some evidence on this question. The claims of my t)ther

W. BoWATER

311

^

« «

CO

K]

OS '^
X

—• s

^ S> =g

o' X m

5' 3 3 ^J -^ S

O -^ -r-

t» 05 a;

p

^ ^ m tq Ei; fij ci;

t-H — * o —

— I oj 2:

2 a . -^ s:

o ^

> .-

cq > =^ i ^ 2 ^■

=q s

j; 6q cq

ft;

S ^ S
M Pm cc

m

■a
a

g o

c

03

a

-I

3 o S

5 o5 2

5 \ S

I I

"^J 1^ ^

I I I

I I

I I 1 1 I

I 2

I I

I 1 1

+

o

^ ^ h h^

312 Hereditji of MeJanhiii in Lep'uloptera

experiment preventing me fmtn giving sufficient attention to the larvae,
and the sj^^^cial liability to disease from which this species suflers in con-
finement, and the usually fatal results which follow inbreeding, militated
against very full results.

I have preserved all the specimens, however, and give a table of
results, as I fail to find a record of even this measure of success by
others in the continuous breeding and inbreeding of this species.

The male parent of family 10.2 was raelanic and is recorded to have
been douhledayaria. The female parent is type. Of the offspring all
the 36 males and many of the females are undoubtedly intermediates.
Five of the females are so nearly black that it is difficult to place them.

The offspring of 7 pairings from this femily were safely reared and
emerged June, 1912 (see table), and one pairing from these was successful,
and the resultant moths appeared June 1913. Pairings have been
again obtained and at the present moment a few descendants survive as
pupae.

It is worthy of note that in the four families, each the offspring of
two intermediate parents, the type sjieciraens total 14 and the melanic
44 (approximately 1:3); and in the four families, each the offspring of one
type and one intermediate parent, the type and the melanic siiecimens
appear in about equal numbers (45 and 48).

1910

Table

Label of

faiuily

10.1

Imagines

Parentage
Male X Female

Type

,- ^ .

Male Female
— 3

Totak
3

Male
2

Intermediate

Female Totals
2 i

Black
Male

Female

*M X T

10.2

— —

—

36

37

73

_

—

T X M

10.5

— 2

2

(32)

(5)

10.3

— —

—

8

1

9

1

4

10.4

1 1

2

7

1

8

—

—

Colchester

—

2

2

1

—

1

Scotland

—

— 1

1

—

—

—

—

—

Totals ... 110
* Jl/= Melanic, but not ueceasaiily true black (doitbledai/aria}.

Apparently, therefore, the M factor is not completely dominant but
seems to give either partial or complete blackening and the cause of
this difference between full melanism and intermediate must be left
open, but one point is clearly shown, namely, that clean Mendelian
segregation occurs (vide 12.2, ll.S, 11.20, 11.29 and 11.34).

W. BOWATER

313

1911 i

1912

inU rO(,/

191

■2 Table.

Label of
family

Imagines

te

Parentage
Male X Female

Type

Intermedia

Male

Female

Totals Male

Female

Totals

; 10.2 / X 10.2 I

11.18

1

4

.■, 1

4

0

10.2 I X 10.2 I

11.20

2

•J

4 10

1-5

2.5

10.2 7 X 10.2 I

11.29

—

2

2 7

o

12

10.2 / X 10.2 I

11.34

.2

1

3 1

1

2

Totals

14

44

10.2 / X ColcL. T

11.2;-,

s

3

11 7

3

10

+

10.2 I X Scot. T

11.38

•5

7

12 9 + '

G + 7

17

10.2 I X Coleh. r

11.41

2

1

3 1

—

1

11.34 T X 11.20/

12.2

9

10

19 14

6

20

Totals

45
Total for 1911
Total for 1912

48

113
40

Total speeimeus shown
: Gyiiandromorphous.

263

Genealogical Table.

1909

1910

Scotland
Type

10.2
Intermediates

Colchester
Type

1911

1912

1913

F,

pupae

13.6 13.7 13.5

314 Hcrcditu of JfcffiiilsiH in Lepido2)tera

It is hoperl that the whnle i(nesti(in can be settled during the next
few years.

In conclusion, with regard to most species which have nielanic forms,
the evidence is not extensive, bnt the weight of evidence up to the
present seems to show that melanism in Lepidoptera frequently follows
the Mendelian Law of Heredity, and in most cases is dominant, but in
some few species is recessive.
' I wish to record my indebtedness to the various observers mentioned,
I'specially Messrs Mansbridge, Prijut, Porritt, J. W. H. Harrison, and
Buckley, for their courtesy in correspondence, and for permission to
include the data of their respective experiments.

DESCRIPTION OF PLATE XXVII.

1 — 12. A Hiphidaays betuUiria.

1. Light type i , Kent.

2. Light type ? , Kent.
i. Dark type g , Kent.

4. Dark type ? , Kent.

5. Litermediate J , Family 11.20.
I.i. Intermediate V , Warwickshire.

7. Intermediate S .

8. Intermediate ? , Family 12.2.

9. Intermediate ^ , Family 11.20.

10. Intermediate 1 , Family 11.20.

11. Var. douhledayaria i ■

12. Var. douhledayivria ? .

13 — IG. Spilosoma lubricipeda.

13. Type 6 , Worcestershire.

14. Var. Yorkshire <f .

15. Var. Intermediate <s .

16. Var. radiata ? .

17 — 19. Boarmia repandaia.

17. Type i , Devon.

18. Banded ? , Devon.

19. Melanic $ , LaucaBhire.

20 — 22. Acidalia rirtiuhiria.

20. Light {ctiitteiierariii) j . Hyeres parents.

21. Dark J, Loudon parents.

22. Melanic ? .

23 — 2.5. Aplectii nebutosti.

23. Type <f .

24. Var. robsaiii s .

25. Var. tliompsoni 2 .

JOURNAL OF GENETICS, VOL. ML NO. 4

■w!ft;% . ■■

'....xi'' '

f^-.

■ 1^:/-^

.-.^ :,^Vi

vV-^/'

- ^ '■ at ; ? *

'■^. .••>

V

,**■"

¥ ^ ^i

.-<''<

-'' '7

PLATE XXVII

^f?rH^^^^\j./.,;?'^^9BL

26

^7

0 >

28

-I-

^9

'^'^my 4c

41

42

'^Ty ' ?;■.

4«

49

W. Bo WATER 315

26 — 33. Odontopera bidentnta. Type forms.

26. Isle of Wight ? .

•27. Family 11.14 ? .

28. Isle of Lewis j .

29. Isle of Lewis <f .

30. Family 11.24 ? .

31. Surrey J .

32. Family 12.64 <f .

33. Family 12.26 <J .

34 — 39. Odontopera bidentata. Melauic forms.

34. Yorkshii-e d .

3.5. Family 11.11 tf . Homozygous parent of 12.34.

36. Family 12.34 g . Heterozj'gous.

37. Family 12.34 9 . Heterozygous.

38. Family 12.11 $, with fulvous tint.

39. Family 12.11 i , with light fulvous patch.

40, 41. Coremia unidentaria.

40. Pieddish banded s .

41. Black banded i .

42, 43. Hemerophila abruptaria.

42. Type ? .

43. Melanic i .

44, 4.5. Abraxas grossulariata.

44. Type ? .

45. Var. varleyata j .

46, 47. Triphaeiiu comes.

46. Type 3.

47. Melanic i .

48, 49. Acidalia contiyiiaria.

48. Type ? .

49. Melanic 3 .

1—12. Selected from 700 specimens in writer's collection.
13, 14. Lent by Mr A. Home.
15—18. Lent by Dr Beckwith Whitehouse.

19, 23, 24, 25. Lent by Mr W. Mansbridge (bred in his experiments).

20, 21. Bred in experiment by Messrs Prout and Bacot, Lent by Professor Poulton.
22. Lent by Mr R. T. Baumann.

26 — 39. Selected from 2100 specimens in writer's collection.

40, 41. Lent by Mr L. B. Prout (bred in his experiment).

42, 43. Lent by Dr Beckwith Whitehouse.

44, 45. Lent by Mr 6. T. Porritt.

46, 47. Bred in experiment by Mr Bacot. Brood B. Lent by Professor Poulton.

48, 49. Lent by Mr W. Buckley (bred in his experiment).

Cnmbriigf :

PRINTED liV JOHN CLAV, M.A.
AT THE UNIVKRSITY PRESS

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clues with him, to lure his curiosity, and to challenge him to thought. The
student will eventually discover that certain periods or writers are more to
his taste than others; he will require, above all, bibliographical guidance.
This he will find in The Cambridge History of English Literature, to which
this Primer may serve as an introduction " — Preface.

Contents. — Book I. Old English Literature to the Norman Conquest. — Book II.
The Middle Ages. — Book III. The Renascence. — Book IV. The Literature of the
Middle Classes. — Book V. The Revival of Romance. — Book VI. The Victorian
Age. — Appendix. — Index.

The Cambridge History of English Literature. Edited by Sir A. W. Ward,
Litt.D., F.B.A., Master of Peterhouse, and A. R. IValler, M.A.,
Peterhouse. Volume X, The Age of Johnson.

Royal 8vo. pp. xvi + 562. Price in buckram, 91. net, in half-morocco,
1 5$. net ; to subscribers 75. 6d. net and 12J. 6d. net respectively.

"As a work of reference," says The Observer, "there can be no doubt
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Some Press Opinions of Volume X.

"This is the most striking volume yet issued of a really great history. It has taken
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"Mr David Nichol Smith's 'Johnson and Boswell ' presents a skilful summary of
Johnson's career and work. Mr Nichol Smith is too sound a scholar to fall into a kind
of smartness currently regarded as paradox, and too much practised in our time. He
makes his points, however, and they are well made. ...The Bibliographies appended to
the present volume continue to prove as serviceable to the student as those in previous
I'olumes of the Cainbridge History." — T/te Westminster Gaaette

COUNTY GEOGRAPHIES

Northumberland. By S. Rennie Haselbunt, M.Sc, F.G.S.

Crown 8vo. pp. xii + iS2. With maps, diagrams and illustrations. Price i;. 6J.
In addition to its natural features, the antiquities ot' the county, in-
cluding the wonderful Roman wall and other remains, are fully described

Bamburgh Castle

and illustrated in the present volume, as are also the churches, castles, famous
men and other interesting features.

For the convenience of tourists and others, the hook is also issued in a smaller
and handier form pjith limp covers and rounded corners at \s. bd. net.

Merionethshire. By A. Morris, F.R.Hist.S.

Crown Svo. pp. x+ i66. With maps, diagrams and illustrations. Price u. 6d.

I-

ySS^'.

^^ ^^*^

The Beach, Llwyngwril

"Both in its verbal descriptions and photographic illustrations, the book does justice
to the most picturesque of Welsh counties." — The Mancliester Guardian

SOUND— AN OLD PROBLEM— BUILDING

Sound. An elementary text-hook for Schools and Colleges. By J. JV. Capstick,

M.A. [Camh.\ D.Sc. {Fict.), Fellow of Trinity College, Cambridge.

Crown Svo. pp. viii + 296. Price 45. 6it.

This new volume in the Cambridge Physical Series is primarily intended
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The author, realising that many of the most ordinary phenomena cannot
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required for the appreciation of the physical connexion between the pheno-
mena described.

The chapter-headings are as follows: Nature of Sound — Elasticity and
Vibrations — Transverse Waves — Longitudinal Waves — Velocity of Longi-
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— Quality of Musical Notes — Organ Pipes — Rods. Plates. Bells —
Acoustical Measurements — The Phonograph, Microphone and Telephone
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Instruments.

A large number of questions taken from Examination Papers set in various
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"Squaring ihe Circle." A History of the Problem. By E. JV. Hohson,
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Demy Svo. pp. viii + 5S. Price y. net.

"The problem which is known to mathematicians as the quadrature
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due partly to its comparative obviousness, partly to its 'proverbial' celebrity
and partly to an exaggerated notion of the gain which would accrue to
mankind by its solution."

Architectural and Building Construction Plates, Part I. Thirty draivings
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are interested in the soil. ..will find this handbook of much interest and value."

73. The Life Story of Insects. By Prof. G. H. Carpenter.

"Prof Carpenter, one of the foremost zoologists of Ireland, has contributed an
excellent primer to the Cambridge Manuals." — 'Dublin Express

74- The Flea. By H. Russell.

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able humour. ..the story is admirably told in language
which is intelligible to the most unscientific reader."

75. Pearls. By Prof W. J. Dakin.

"An interesting and simply written account of
the most important facts about pearls, pearl fishing
and pearl formation." — The Athenteum

• 76. Na'val Warfare. By J. R. Thursfield,

M.A. IVith an Introduction by Sir

Charles Ottley.

"This little book," says The Daily Telegraph, "has

a mission. It is the quintessence of the naval wisdom

of the ages and. ..will achieve no small end if it falls

into the hands of the public men of the Empire."

77. The Beautiful. By Vernon Lee.
"An excellent and lucid discussion of aesthetics

fascinatingly put before the reader." — Church Family
Neivspaper

78. The Peoples of India. By J. D. An-

derson, M.A.

" Mr Anderson is an ex-Indian civilian of large
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terly summary of a wide and complex subject." — The
Guardian

79. The Evolution of Ne<w Japan. By Prof J. H. Longford.

"The author was for many years British Consul at Nagasaki and his work is a
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not easy to procure." — The Guardian

80. A Grammar of English Heraldry. By IV. H. St John Hope, Litt.D.

"Dr St John Hope has done nothing better than his Manual One sees that heraldry

is not a musty topic of no earthly value but a subject that is a valuable handmaid of
history and art." — Pall Mall Gazette

A full list of the series will be sent on application. For particulars of
the special sets in cases see p. ii oj wrapper.

9 1—5

(Froni Hope's Grammar
En^liih Heraldry.^

»/

RUBBER PLANTING— FLIES AND DISEASE

Rubber and Rubber Planting. By R. H. Lock, ScD., Inspector H.M.
Board of Agriculture and Fhheries, sometime Assistant Director of Botanic
Gardens, Ceylon.

Crown 8vo, pp. xiii + 245. With lo plates and 18 text-figures.
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To make the book suitable to as wide a circle of readers as possible the
author has aimed at combining an accurate account of the scientific side of
rubber planting with a certain amount of practical information for the
benefit of the prospective planter.

"Each subject is dealt with concisely and the facts presented in an interesting and
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any point." — The Financial Times

"...Should meet with a wide welcome. The authoritative account given by
Dr Lock is not only extremely interesting to read, but should also be of real value to
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rubber company The description of the industry gains much by the illustrations,

which include. ..photographs. ..and excellent line-drawings, specially made." — Knonv-
leJge

Flies in relation to Disease. Non-Bloodsucking Flies. By G. S. Graham-
Smith, M.D., University Lecturer in Hygiene, Cambridge.

Demy Svo. pp. xiv -1-292. With 24 plates and 32 text-figures.
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Press Opinions oj the first volume of the Cambridge Public Health Series.

"In this book Dr Graham-Smith has collected the most important and reliable
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"This is just the book for students who either are, or are to be, occupied with
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10

BRITISH PLANTS— SCHOOL GARDENING & HANDWORK

Genera of British Plants arranged according to Engler''s Syllabus der
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tomed in the British Floras that have hitherto appeared, and to serve as an

introduction to the use
and the Pfan-zenreich.

Educational School Gardening and Handivork. By G.
F.R.H.S., Inspector of Educational School Gardening,
Education Co)nmittee. IVith an Introduction by the
Hothouse, Chairman of the Somerset County Council.

of the Syllabus, the Naturliche Pflatrzenfamilit

IV. S. Brewer,

Somerset County

Rt Hon. Henry

Crown Svo. pp. xii + 192. With 75 illustrations. Price zs. 6d. net.

li

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12

Books puhlished since 6 November- 1 9 1

THEOLOGY

Forgi'veness and Suffering, ^ Study oj Christian Belief. By Douglas
White, M.D. See p. 3. '

St Basil ihe Great. A Stuety in Monasticism. By IP^. K. Lowther Clarke.
See p. 3.

The Parables of the Gospels in the Light of Modern Criticism, Huhean

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The Story of Ahikar, From the Aramaic, Syriac, Arabic, Armenian,
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LATHS!

Linrn, Book XXVII, Edited by S. G. Campbell, M.A., Fellow and Classical
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13

ENGLISH LITERATURE— FRENCH— GERMAN

ENGLISH LITERATURE

A Primer of English Literature. By IV. T. Young, M.A. See p. 6.

A Book of English Prose. By Percy Lubbock, M.A., King's College,
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Part I contains about 30 short extracts, ot about tour pages each, from the most
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Part II contains a fuller selection ot similar passages. In each part the notes are
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Essays on Social and Political Questions. By J. Howard IFhitehouse,
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Charles Lamb : Essays of Elia and The Last Essays of Elia. Edited
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Each volume contains an introduction, notes and indexes. The editor acknow-
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A Handbook of Precis- Writing. JFith Graduated Exercises. By E. Derry
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14

CLASSICAL HISTORY— MATHEMATICS AND PHYSICS

CLASSICAL HISTORY AND LITERATURE

Essays and Studies Presented to William Ridgeivay, Sc.D. Edited by

E. C. Quiggin, M.A. See p. 2.
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HISTORY

The Puritans in Po'wer. By G. B. Tatham, M.A. See p. 5.
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Bartotus of Sassoferrato, His Position in the History of Medieval
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International Lanv. Part IL War. By John IVestlake, LL.D., K.C.,

late JFhewell Professor of International Law in the University of
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MATHEMATICS AND PHYSICS

'* Squaring the Circle." By E. TV. Hobson, Sc.D. See p. 8.

Sound. By J. TV. Caps tick, M.A. See p. 8.

An Algebra for Preparatory Schools. By Trevor Dennis, M.A., Mathe-
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15

GEOGRAPHY & TRAVEL— PSYCHOLOGY— EDUCATION

GEOGRAPHY AND TRAVEL

An Elementary Commercia.1 Geography. By H. R. Mill, D.Sc Revhed
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The Madras Presidency). By E. Thurston. See p. 4.
Northumberland. By S. R. Haselhurst. See p. 7.
Merionethshire. By A. Morris. See p. 7.

ARCHAEOLOGY

The Place-Names of Nottinghamshire, their Origin and Development.

By Heinrich Mutschmann, M.A. {Liverpool), Ph.D. [Bonn), Lecturer
in German and in Phonetics at the University College, Nottingham.
Cambridge Archaeological and Ethnological Series. Demy 8vo. pp. xvi
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PSYCHOLOGY

Obliviscence and Reminiscence. By Philip Boswood Ballard, M.A. The
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EDUCATION

A National System of Education. By J. H. IT hit chouse. See p. i.

The Purpose of Education. An Examination of the Education Problem in
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16

BOTANY— CAMBRIDGE UNIVERSITY— JOURNALS

BOTANY

Genera of British Plants. By H. G. Carter, M.B. See p. 1 1 .

School Gardening and Handwork. By G. IF. S. Breiver. See p. 1 1 .

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Antionncerufnts

CAMBRIDGE UNIFERSITT CALENDAR

In 1796 appeared the first Cambridge University Calendar. It was
a small volume of 190 pages, edited by G. Mackenzie, B.A., of Trinity,
and contained " a list of the present members, the livings in the gift of
each College, with their incumbents; some useful particulars concerning
the Fellowships, Scholarships etc. Professors, Officers, Prizemen etc."
It was printed by Benjamin Flower and published by W. Page, and in
the Preface a pious hope was expressed that it would be " neither useless
nor uninteresting to the Members of the University." A note on the
Origin of the University, characteristic of the age of faith, occupied
the first page :

"The origin oi Cambridge as an Uniuersity is very doubtful. We are however

informed that one Cantaber, a Spaniard, about 370 years before Christ, is intitled

to this honour ; Certain it is, that after many years laying desolate, Sigebert,

King of the East Angles, restored it a.d. 630."

Except tor the year 1798, the Calendar has appeared annually smce
1796. In 1 803, the title was temporarily changed to The Cambridge
University Register, and the following year's issue, edited by "a member
of the Senate," was dedicated to William Pitt and the Earl of Euston.

It was printed at the University Press iti 1803 and 1804, and this has
also been the case from 1826 to the present time.

With its publication, the name of Deighton has been associated since
1803, ^"'' '"^^ proprietorship has recently been transferred from the present
firm of Messrs Deighton, Bell & Co. to the Syndics of the Press, wlio
will be responsible for its publication, in a new format, in 1914 and
subsequent years.

CAMBRIDGE MANUALS

Five new volumes will be added to this series at an early date.

These will include two scientific volumes on The San and Coal-mining
by Professor R. A. Sampson of Edinburgh and Mr T. C. Cantrill of the
Geological Survey respectively ; one on a pressing industrial problem
entitled Economics and Syndicalism by Professor A. W. Kirkaldy of
Birmingham; a short history of The Royal Navy by Mr John Leyland ;
and a volume of lectures on Military History (originally given at Trinity
College) by the Hon. J. W. Fortescue.

CAMBRIDGE BRITISH FLORA

The Syndics of the Press have pleasure in announcing their arrange-
ments for the issue of a new, comprehensive and fully illustrated British
Flora. The work will be completed in about ten volumes, which, so far as is
practicable, will be issued annually. Publication of the work will begin
with the issue of Volume II, which will be ready in March 1914.

18

CAMBRIDGE BRITISH FLORA

The styles of binding and the prices will be as follows ; —

PublisheJ Price Price per 'volume

per volume to subscribers to

the njjlwle ^vork
Paper boards, with canvas back and paper label
each volume in two parts, the first containing
the text and the second the plates ... ... £z. \os. net £2. 5/. net

Quarter morocco, in two parts divided as above ^6 net £^. ^s. net

Paper boards, with canvas back and paper label,
in one volume the plates mounted on guards

and bound interspersed with the text ... ^^3 net £1. 1 55. net

Quarter morocco, in one I'olume the plates
mounted on guards and bound interspersed
with the text ... ... ... ... £6 net £$■ 5'- «f'

Each indigenous species of flowering plants, many naturalised species
and many varieties and formae will be illustrated from the pen-and-
ink drawings of British flowering plants which have recently been
presented to the University of Cambridge by Mr E. W. Hunnybun.
Each plant or portion selected has been drawn natural size, and will be
reproduced without reduction or enlargement. In addition to the main
drawing of each plant, there are also enlarged drawings of critical organs.
Each drawing has been made by Mr Hunnybun from a fresh plant, the
name of which has been vouched for by some competent authority whose
letter of identification is preserved in the Cambridge University Herbarium.
Such a set of drawings is quite unique in the history of botany. The high
artistic merit and scientific value of tlie drawings are admitted by all who
have seen them.

The Flora will be written by Dr C. E. Moss, Curator of the Cam-
bridge University Herbarium. Engler's system of classification will,
generally speaking, be followed. This system is becoming very generally
adopted : already there are German, Swiss, American and other Floras
based upon it.

The systematic descriptions will be in English, not in Latin ; and
the geographical distribution of important groups will be fully stated.
Distribution-maps will be freely provided.

The critical elucidation of the more difficult genera will receive
careful attention. A large number of botanists have kindly promised
their aid, and, in order to keep a uniform standard of treatment, Dr Moss
will act as Editor of the genera treated by the various specialists.

Volume II will deal with the earlier Dicotyledonous families, and thus
will include most British trees, such as Poplars, Willows, Birches, Oaks and
Elms, as well as Docks, Goosefoots and Glassworts. It will contain an
important contribution by the Rev. E. S. Marshall on Birches.

19

FORTHCOMING BOOKS

FORTHCOMING BOOKS

Among other books which will be ready immediately is the first volume
of Mr J. H. Wylie's History of the Reign of Henry V. This work, which
will probably be completed in 4 volumes, will be a companion to the same
author's Hhtor\ of Henry IV^ published by Messrs Longman in 1898.

Sir Charles Waldstein's lectures on Greek Sculpture and Modern Art,
delivered before the Royal Academy, will be published in book form with
the addition of more than 70 full-page plates illustrating ancient and
modern sculpture.

The last course of lectures given by the late Dr Verrall as King
Edward VII Professor of English Literature have been edited by Mrs
Verrall and will appear under the title Lectures on Dryden.

The Naval and Military Serifs will be inaugurated by a volume on
Ocean Trade and Shipping by Mr Douglas Owen and a collection of
Naval and Military Essays, being papers read at the International His-
torical Congress in 1913.

To the Handbooks of Liturgical Study will be added Mr T. Thompson's
volume Jhe Offices of Baptism and Confirmation.

Harrington and his Oceana is the title of a historical study by Mr
H. F. Russell-Smith, in which he examines the political theories of
James Harrington with special reference to their connection with the
American constitution.

The second part of Dr E. C. Clark's History of Roman Private Law
will be published in two volumes.

Dr E. G. King's The Poem of Job is an English translation in the
metre of the original poem.

Knoiv Your Own Mind is described by the author, Mr W. Glover,
as " a little book of practical psychology," and seeks to interest the man
in the street in the study of conduct.

Mr S. A. McDowall's Evolution and the Need of Atonement, of which
a second edition will be ready immediately, has been revised and con-
siderably enlarged.

A volume on Photo- Electricity by Dr A. LI. Hughes of the Rice
Institute, Texas, will be added to the Cambridge Physical Series.

The Respiratory Function of the Blood is a medical treatise b)-
Mr Joseph Barcroft.

Mr S. Holmes has written a linguistic study of Joshua, in which the
Hebrew and Greek texts are compared.

Dynamics, by Professor Horace Lamb, will be a companion volume to
the same author's Statics published about a year ago.

To the French volumes in the Pitt Press Series will be added an edition
of La Jeunesse de Cyrano by Mr H. A. Jackson of Winchester College
and to the Latin an edition of the Phormio by Mr John Sargeaunt pf
Westminster.

20

Carpentum of Agrippina the elder

Companion to Latin Studies. Edited by Sir j. e.

Sandys, Litt.D., F.B.A. Second Edition. Royal 8vo. With 2 maps,
141 illustrations, and four indexes. 18^. net.

"In the single volume before
"" us it is really possible for the

first time to obtain a conspectus
of almost all that is definitely
known about Roman environ-
ment, life, and thought — The
Book is a thesaurus of sane
learning in a readable form.
Varro or Pliny or St Isidore of
Seville would have studied it
with a growing wonder and en-
lightenment ; for not Rome only,
but the history of all knowledge
about Rome, is here recalled to
its first beginnings." — Times

"Every subject which is in
any wayconnected with the study
of Italy, of its history, its people,
its laws, its art, and its literature
is here discussed by some com-
petent authority. The informa-
tion given is up to date, and is
offered for the most part as fully
as the natural limits of so com-
prehensive a work allow.. ..As a
book of reference the volume will
be of the highest value." —

Westminster Gazette

"This volume is a complete
cyclopaedia of Roman studies ;
and in nearly 900 pages and half
a million words contains the
carefully adjusted result of re-
cent inquiries into every depart-
ment of Latin lore. It is, in a
very remarkable degree, accu-
rate, complete and abreast of
modern discovery ; and we con-
gratulate the University, the
contributors, and the editor on
the signal success of an ambitious
project." — Saturday Review

Private portrait (age of Gallienus)

Quantity and Accent in the Pronunciation of Latin. By

F. W. WeSTAWAY. Crown 8vo. pp. xvi -(- 1 1 2. 3^-. net.
"This book," says The Athenaeum, "should be read and kept for
reference by all teachers of Latin."

The author makes a spirited attack upon "the remnant of the old school"
which still clings to its "duU'sy dome'um " {dtilce domiiin) and its "nice-eye
pry-us" {iiisi priUs) and addresses his book (i) to private students who desire
to learn to pronounce and to read Latin correctly, and (2) to those who feel
that their acquired pronunciation needs overhauling.

MENDEL'S PRINCIPLES
OF HEREDITY

By W. BATESON, M.A., F.R.S., V.M.H., Director of the John Innes
Horticultural Institution. Third impression with additions. With
3 portraits, 6 coloured plates, and 38 other illustrations. Royal 8vo.
12s. net.

In the past three years the progress of Mendelian analysis has been very
rapid, and the author has endeavoured in a series of brief Appendixes to
acquaint the reader with the nature of the principal advances made.

"This third edition shows how rapid has been the progress of Mendelian analysis. It deals
with heredity of colour, heredity and sex, gametic coupling and spurious allomorphism, double
flowers, evidence as to MendeUan inheritance in man, biological conceptions in the light of
Mendelian discoveries and practical application of these principles. An appendix contains a
biographical notice of Mendel and translation of his papers on hybridization and hieracium.
The work is beautifully illustrated." — Chicago Medical Journal

"A new impression cannot fail to be welcomed Mendel's Principles of Heredity is already

a classic. It marks a position of stability towards which previous work is now seen to have
logically converged, and from which new and active research is to-day no less logically diverging.
The various waves of biological thought are constantly intersecting, mingling, and passing on
with altered rhythm, but it rarely happens that so many meet together at a nodal point as during
the last decade. ...As an analysis of that point, as a picture of how it has come into being, and as
a foreshadowing of happenings in the near future, Mendel's Principles stands alone, and it is
good to know that the generation of students now growing up cannot be cut off from the posses-
sion of a book so full of inspiration." — Gardeners' Chronicle

THE METHODS AND SCOPE
OF GENETICS

By W. BATESON, M.A., F.R.S., V.M.H.
Crown Svo. Is. 6d. net.

" Professor Bateson tells how Mendel's law works out with the colours of certain flowers,
moths, and canaries, and with colour-blindness in men and women. More than this, he describes
the outlook over this field of research in a manner that will greatly interest and attract aU in-
telligent people, for, as he rightly says, ' Mendel's clue has shown the way into a realm of nature
which for surprising novelty and adventure is hardly to be excelled.' " — Morning Post

Cambridge University Press, Fetter Lane, London
C. F. Clay, Manager

JUST PUBLISHED

Crown Svo. 3s. 6d. net

CONTROLLED NATURAL SELECTION AND
VALUE MARKING

By J. C. MOTTKAM, M.B. Lond.
LONGMANS, GREEN & CO., 39 Paternoster Row, London, E.G.

CONTENTS

All Rights reserved

PAQB

J. W. H. Harrison and L. Doncaster. On Hybrids between Moths
of the Geometrid Sub-Family Bistoninae, with an Account of the
Behaviour of the Chromosomes in Gametogenesis in Lycia (Bistori)
Hirtaria, Ithysia (Nyssia) zonaria and in their Hybrids. (With
Plates XVII and XVIII) 229

Onera a. Merritt Hawkes. On the Relative Lengths of the First
and Second Toes of the Human Foot, from the Point of View of
Occurrence, Anatomy and Heredity. (With Plates XIX — XXI
and 6 Text Figures) 249

Rose Haig Thomas. The Transmission of Secondary Sexual
Characters in Pheasants. (With Plates XXII— XXVI and
2 Text Figures) 275

W. BowATER. Heredity of Melanism in Lepidoptera. (With Plate

XXVII) 299

The Journal of Genetics is a periodical for the publication of records of original
research in Heredity, Variation and allied subjects. The Journal will also, from time to
time, contain articles summarising the existing state of knowledge in the various branches
of Genetics, but reviews and abstracts of work published elsewhere will not, as a rule, be
included. Adequate illustration will be provided, and, where the subject matter demands
it, free use will be made of coloured plates. The Journal will be issued in parts as material
accumulates, and a volume, appearing, so far as possible, annually, will consist of four such
parts.

Volumes 1 and II (1910 — 13) are now ready. Price in four parts, paper covers, 30s. net
each ; bound in buckram, 34s. 6d. net each. Buckram binding cases can be supplied at
2«. 6d. net each. Subscribers' sets can be bound for 4s. 6d. net per volume, including the
case.

Papers for publication may be sent either to Mr Bateson, Manor House, Merton
Park, Surrey, or to Professor Punnett, Whittingehamc Lodge, Cambridge. Other
communications should be addressed to the University Press, Cambridge.

Papers forwarded to the Editors for publication are understood to be offered to the
Journal of Oenetics alone, unless the contrary is stated.

Contributors will receive 50 separate copies of their papers free. A further 50 may be
had at cost price : these should be ordered when the final proof is returned.

The Subscription price for a volume is 30s. net (post free) payable in advance ; separate
parts 10s. each net. Subscriptions may be sent to any bookseller, or to Mr C. F. Clat,
Manager, Cambridge University Press, Fetter Lane, London, E.C.

The Cambridge University Press has appointed the University of Chicago Press agents
for the sale of the Journal of Getietics in the United States of America, and has authorised
them to charge the following prices : — Annual subscription, f 7.50 net ; single copies
?2.50 net each.

CAMBRIDGE: PBINTED BY JOHN CLAY, M.A. AT THE UNIVERSITY PRESS

New York Botanical Garden Libran

3 5185 00266 2052

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