See a SR a “het tea yy AD Ay THIS eat aber 3 +a Pus Mendel Journal October 1909 — 1. ORIGINAL RESEARCH. Parthenogenesis in Nicotiana -. Mrs. R. HAIG THOMAS 2. MENDEL COLLECTION OF HUMAN PEDIGREES. Inheritance of Suicidal Mania Pedigree of Tuberculosis Pedigree of Skin Colour in Human Hybrids. A case of Segregation of White Skin in a Quadroon Fraternity 3. PAPERS READ TO THE MENDEL SOCIETY. The Evolution of Man - - J. T, CUNNINGHAM Plea for operation of more virile sentiment in Human Affairs : - nie AGP OMUDGE Mendelism and Sex - : : - ~~. . €. €. HURST 4. METHODS AND RESULTS - “ARDENT MENDELIAN” Present position of Mendelians and Biometricians Skin Colour in Human Hybrids : Variation in the single combs of Fowls 5. MISCELLANEA. Science and Democracy Nature of Scientific Hypotheses 6. REVIEWS OF BOOKS. Copyright All Rights Reserved Printed and Published for the Mendel Society by TAYLOR, GARNETT, EVANS, & Co., Ltd., 54, Fleet Street, London, E.C., and Manchester No. oe Price 2/6 nett f ee % ae AI I> BOTANE Mendel 4 ournal PROLOGUE. Mendelism is a subject which has come to stay and to play an important part in human affairs. In Agriculture, in Horticulture, in the Prize Pens, and in Sociology, its voice will be heard. It will not be as a voice in the desert, but as a world-vibratory one, uttering its pronouncements, admonitions, and definitive conclusions, based on the solid and unshake- able ground of accurate experiment, wherever culture and life come into contact. A subject like Mendelism, full of complexities, and hable to be misunderstood by the careless, the hasty, and the unwary, necessarily runs many dangers among the shoals of human affairs. There are the dilet- tante philanthropists and scientists, and social refor- mers; and there are newspaper writers possibly seeking to ingratiate themselves with their numerous readers, by assuming the garb of “ gods who destroy false idols,” which they themselves first set up. Not Mendelism, nor its progress, but the application of it to human affairs, is endangered by them. A great subject like this; one with not only an academic importance, but clearly of great practical importance also, and having a bearing upon some of the greatest phases of human activities, so far as 2 THE MENDEL JOURNAL its presentation and promulgation are concerned, has two courses open to it. These two alternatives are inevitable. Mendelism differs in this respect from subjects which are purely academic, because these are only presented and discussed in scientific societies and in their journals. But Mendelism is a subject which comes into the category of the ““ Humanities ” ; it will be discussed in various circles, by people more or less well acquainted with it, and more or less antagonistic to it. Must its wider promulgation necessarily be left wholly to them? We think all will agree that a better alternative lies before us. It is better that Mendelism shall be presented to a wider public by men who believe in its truth, who foresee its future, and who recognise their responsi- bilities in the work they do. That is one of the objects with which the “‘ Mendel Journal” starts upon its career, and which forms the far distant beacon-light towards which it will consistently steer. But it has another object. It is to gather for the Science of Genetics a harvest rich in facts relating to human pedigrees and the inheritance of normal characters as well as of peculiarities. To find these the seeker must quit the experimental garden and the cloister, and he must pass out into the world of his fellows. From them shall the grain be gathered in order that it may be garnered in these pages. A golden field, ripe for the harvest, awaits the coming of the Mendelian reapers. We appeal to all who are acquainted with families in which peculiarities and markedly contrasted normal characters, have run PROLOGUE 3 from one generation to another, to send details to the Editor of this Journal. Such contributions will be treated in the strictest confidence, they need not necessarily be published, and will not be published without the consent of all who are concerned, and then only in a form approved by them. While it is clear that medical men have many unique opportu- nities for acquiring knowledge of pedigrees of this kind, it is hoped that contributions from laymen will also be forthcoming. In future numbers of this Journal, prominence will be given to matters pertaining to agricultural and horticultural practices and problems. It is also contemplated to make it a medium by which authoritative advice and direction may be given in the form of answers to questions upon matters of general interest, relating to problems of cattle, cereal, and plant breeding. In this way, it is hoped that the Journal may become a medium of great value to all who are engaged in the breeding of live stock of all kinds, and to those who are concerned in the production and fixing of new varieties of flowers, or of leguminous and cereal stocks. It is certain that much money and time have been wasted in the past, owing to the haphazard methods and erroneous ideas which were employed in agricultural and horti- cultural practices. It is perhaps not too much to anticipate that this Journal may at least be more valuable than a Royal Commission and as competent as a Government Department, to advance the scientific treatment of 4 THE MENDEL JOURNAL matters of the greatest commercial importance to the country. Any questions directed to the Editor, bearing upon the scientific breeding of cattle and plants, will, as far as possible, receive full consideration. in the next number of the Journal. Business communicatiens should be addressed to the Manager, Office of the Mendel Journal, ¢/o Messrs. Taylor, Garnett, Evans, & Co., Ltd., 54, Fleet Street, E.C. Literary contributions, or questions relating to the scientific breeding of cattle and plants, or pedigrees of human families, should be addressed to the General Editor. ORIGINAL RESEARCH. (1) Parthenogenesis in Nicotiana. By ROSE HAIG THOMAS. I NoTIcED last year and this summer that in several different cross fertilisations, made between different species of Nicotiana, some plants bearing only the characters of the ovule parent were produced along with others which were apparently Fi hybrids. Upon the first and second occasion when I obtained these results—which occurred with reciprocal crosses of Nic. Sylvestris x Nic Affinis.—I thought they must be due to accident, for muslin bags had been used, and it was possible that pollen from adjacent flowers of the same plant may have gained access through the bag. I repeated the experiment a third and a fourth time, with every precaution, such as the use of wax-paper bags and the cleansing of the pollinating instruments in spirit before use. At both repeti- tions of the experiments the same phenomenon was manifested. I then began to suspect the cause, and determined to test Nicotiana for parthenogenesis. In the first few trials the method employed was to cut off the anthers alone, but later, after success attended these I cut off both stigma and 6 THE MENDEL JOURNAL anthers; the result remained the same, the buds developed into full bloom, the corollas withered, the capsules set, the seed in due time ripened, and if left on the plant long enough the capsules split open and shed their seed. My first trial for parthenogenesis was with Nie. tabaccum Cuba, raised from some seed I gathered from a plant in the well-known garden of Casa Loring, near Malaga. The gardener there told me that this plant had grown from seed brought over from Cuba, and that it had been gathered from the finest tobacco plants grown in the island. This plant is taller than other N. tabaccums, and is 6Ht. to 7ft. in height, and the stems are very thick; it flowers at first in a terminal cluster and afterwards axially. The limb and tube of the corolla are pure white; the corolla is sometimes four petalled with four stamens, some- times five petalled with five stamens, and both forms are found on the same plant. It is a freely self-pollin- ating plant, for under protection from insects it will seed every blossom. On July 15th, 1909, I cut off all the anthers from five young green buds on a spray of Nie. tab. Cuba, and covered them with a wax-paper bag, which was wired on in the usual way. At the same time all the other buds and blossoms on the spray were removed. On July 24th the spray was uncovered and it was found that only one ovary had failed, the other four capsules having set seed. One or two tiny buds were sprouting; these were pinched off, and the bag replaced over the seed. I at once started a PARTHENOGENESIS 7 second experiment on the same plant to confirm this first one, and proceeded in exactly the same manner. On the same day I experimented with another plant, using the same precautions. This plant was a hybrid of Fi N. YLabaccum Cuba x N. Fi P.* (N. Sylvestris x N. Sandere x N. affinis Fi R.*) On August Ist, the eighth day alter, all the capsules were found set on both these plants. I now deter- mined to test every Nicotiana species and variety and hybrid flowering in my greenhouse and garden, and succeeded in setting parthenogenetic seed on the following :— Number of Successful Species. Experiments. Notes. Nic. Suavolens ...| Lspray onl plant. | = Nic. Sylvestris ...| 3 sprays on 3 plants. — Nic. Sanderze ...| 2capsules on 2 plants. | — Nic. Tabaccum Cuba..| 5 sprays on 2 plants. | | Parthenogenetic seed sowed 9th Sept., germinated on the z1st Sept. Nic. Tab. Mirodato .... 1 spray onl plant. | Asia Minor; seed obtained from the Board of Trade. F2 Nic. Sylvestris x Nie. Affinis ...| 2 capsules on | plant.) This parthenogenetic seed sowed 4th Sept., germinated : : 12th Sept. Fi Nic. Sylvestris x Nic. Tab. Cuba ...| 2 sprays on 2 plants. —- Nic. Tab. Cuba x Fi P.* (Sylvestris x Fi R.* | Sander x Affinis)...| 3 sprays on 3 plants. = Nic. F2P.* (Sylvestris x Fi R.* Sandere x Aftinis) es ...| 4 sprays on 4 plants. == Nic. Fi P.* Sanderze Affinis aoe ...| 1 capsule on 1 plant. — fo lnee—snede Pe — purple: 8 THE MENDEL JOURNAL The first parthenogenetic seed I examined was that of Nic. Tab. Cuba; half the seed was round, full, sound looking; the other half flattened, poor, little likely to germinate. The parthenogenetic seed was compared with selfed seed of the same variety ; in this latter nine-tenths was full, sound seed. It seems possible that in this variety all the ovules are not capable of parthenogenesis, and that those not set might have proved fertile to a pollination either self or cross. But in the hybrid Nic. Sylvestris x Nic. afinis F2 the parthenogenetic seed was all round, full, and sound. In order to ascertain the condition of the pollen in the Nicotiana buds, within about twenty-four hours of expansion, I gathered from Nic. Sylvestris = Nic. Tab. Cuba Fian unopened bud 23in. in length, and split it open; the style was two-eighths of an inch longer than the stamens, and the anthers were green and solid. I cut one and placed the section under the microscope; the pollen was miniature, and floated in a colourless liquid that dried up after a quarter of a minute’s exposure to the air. As the anthers dehisce at the opening of the corolla, the development of the pollen is probably very rapid towards the end. The observation of the immature condition of Nicotiana pollen in the young bud at a time subsequent ‘to that chosen forthe removal ofthe anthers and stigma testifies both to the ease with which, in this plant, trials for parthenogenesis can be made, and to the reliability of the experiments. PARTHENOGENESIS 9 The cutting off of stigma and anthers in the earliest stage of the young bud does not in the least affect its development; the tube grows to its full length, the corolla expands to its full width before withering, and there is no more delay in setting the capsule than under normal selfing occurring on other sprays of the plant. In these attempts to obtain parthenogenetic seed, failures resulted in some of the hybrids, but these were not more numerous than the failures from attempts to self the same hybrids. The fact remains that parthenogenesis was dis- covered in ten species, varieties and hybrids of Nicotiana, and it is possible wil! be found in all of them if the right period is chosen for the trial, ¢.e., when the plant is beginning to go off its fullest bloom. Amongst the hybrids I found success more likely to attend a test made at this stage, and also when that test was made on a spray which had already seeded one or two of the lower blossoms. Amongst the species and varieties this is not so necessary. In the Tabaccums success was unfailing. When we remember the wide geographical dis- tribution of Nicotiana, the occurrence of partheno- genesis in an Australian plant, like NV. Suavolens, in the Tabaccum varieties,and in South American species, points to the conclusion that it is an ancient character in Nicotiana, and had developed in this genus before the separation of the possible land connection between Australia and South America. Or, if that conclusion is questioned, we are led to infer that partheno- genesis has arisen independently in different species of the genus. 10 THE MENDEL JOURNAL An important consideration which this dis- covery of parthenogenesis in Nicotiana opens to discussion, is the supposed hybrid nature of UN. Sanderw. This plant is said to be a hybrid of N. Forgetiana =x N. affinis. The former species was brought by one of Mr. Sander’s travellers from South America, and it is said was then crossed with UN. affinis. But the true history of this hybrid appears to be shrouded in mystery. In the light of these observations on partheno- genesis in Nicotiana, it is conceivable that Sander may have obtained amongst the F: offspring from his cross of N. Forgetiana « N. affinis not only some true hybrids which he possibly destroyed, but’also some individuals derived from the parthenogenetic seed of N . Forgetiana. He may have selected these and sent out the seed as the type of the supposed hybrid NV. Sanderw. In other words, N. Sanderw may simply be N. Forgetiana. If this is so, it will explain what is, from the Mendelian standpoint, the remarkable fact that the supposed hybrid N. Sanderw breeds true to seed, for this is a phenomenon which is not expected to occur, on Men- delian principles of gametic purity and segregation. Parthenogenesis in Nicotiana perhaps explains a fact which I have frequently observed, namely, when Nic. Sanderw is growing alongside of other varieties of Nicotiana in the open it always seedstrue. This remark applies equally well to other Nicotiana which I have grown. THE MENDELIAN COLLECTION OF HUMAN PEDIGREES.” (1) Inheritance of Suicidal Mania. The Families of Aand B. Pedigree Chart I. BY GEO. P.. MUDGE. I am indebted to Miss Gertrude Flumertelt, who was recently one of my students at the London School of Medicine for Women, for the facts of this pedigree. I cannot sufficiently express my obligation to her for the interest which she manifested in this case, and for the trouble and care which she took in elucidating the various facts. THe Facrs OF THE PEDIGREE. This pedigree was constructed in 1907, by enquiry of some intimate friends of the Bfamily.t There are two families concerned, and they have apparently lived for many generations in an English village. Both families are very respectable, and their members as a whole are moderately well off, while some members have been wealthy farmers. As already indicated, two families are primarily involved in the pedigree, namely, the A and B families. There exists a tradition in the village in 4 * The Editor will be glad to receive from medical men and others, family pedigrees showing the transmission of disease, peculiarities, abnormalities, or of marked abilities, but not necessarily for pubtica- tion. See Prologue. +The symbolic letters employed to indicate the two families concerned do not in any way reveal their identity. The two first letters of the alphabet have been chosen for the necessary purposes of description. 12 THE MENDEL JOURNAL which they live that death by means of self-shooting belonged primarily to the B family, and death by self-drowning to the A family. The two families have intermarried, and among their descendants three forms of suicide are manifested, namely, the two original forms, by shooting and drowning, and a new form, by taking poison. Before the present history commences, according to tradition, the two families were unrelated. One peculiar and singularly significant feature of the case is the very characteristic mode of drowning adopted by the victims of the suicidal mania. It seemingly indicates that the manifestation of the mania is not a matter of opportunity or suggestion, but is an inherent impulse, which all external influ- ences are powerless to check or destroy. It 1s indi- cative of an innate determination so pronounced that it would call forth our admiration were it manifested in nobler ways. The victims proceed to drown themselves by lying down, and then forcing their faces into a pool of water only a few inches in depth until death results. Plate I.—‘* A Pedigree of Suicidal Mania.” C. M. = Cousin marriage. Suicide by drowning. Has been temporarily placed in asylum. Still living in year 1907. = Suicide by taking poison. ah ae shooting. ie areatal generation. Tey , P., P,., P,., and P,., = The different generations. The eae Sue imeicabe those members who have manifested suicidal mania or whose behaviour has caused their friends anxiety. The ringed symbols indicate the normal persons. The ringed symbols marked with a transverse line represent unknown persons, The numbers are merely for descriptive purposes. i) ll eee eRe ll || Pedigree Chart I. The ‘‘ Drily. 2 elt eee ae re P; Pedigree Chart I. The ‘‘Shooting” or B Family. The ‘‘ Drowning ”or A Family. “D2- HUMAN PEDIGREES 13 The tendency to manifest the suicidal mania developes, as a rule, between twenty and twenty-five years of age. There is one marked exception to the tule. It is the male member, No. 6 in the P generation. (Pedigree Chart 1.) He was a wealthy farmer, and was very much respected. He had apparently lived in his native village all his life. Notwith- standing that he had been very successful in his affairs, at fifty years of age,in the July of 1907, he committed suicide by shooting himself. An announcement of his death, accompanied by an obituary, appeared at the time in several of the weekly newspapers. It appears there was no reason for the suicide. But it is significant that through- out his manhood, according to the account of his relatives supplied to us, he was regarded as “ crazy,” though he was never confined in an asylum. As is usual in constructing a pedigree of this sort, one finds that a few individuals are unknown, or the memory of them is lost, and we are perforce compelled to supply their place by hypothetical persons. These persons, however, we know must have existed. There are five of them in the present pedigree, and they are indicated by the cross line passing across the symbol which stands for them. They are Nos, 1-4 in the P, generation, and No. 2 in the P, generation. In the P, generation it is known that other members besides those shown in the pedigree existed, but all family account of them is lost, and I have not yet been able to trace them. 14 THE MENDEL JOURNAL DETAILS OF THE INDIVIDUAL MEMBERS. P., GENERATION. The mode of death of the four individuals repre- sented is unknown. The existence of these four members is postulated. The family tradition is that the wife of Mr. A No. 1, and of Mr. B No. 3, were normal, and that the swicidal mania came down through the two husbands of this generation. P, GENERATION. In this generation Mr. A No. 1, and Mr. B No. 3, are known to have been brothers-in-law. Since Mr. B No. 3 was known to have married a woman No. 4, unrelated to either family, it follows that Mr. A No. 1 could only be brother-in-law to Mr. B No. 3 by marrying a sister of his. Accordingly, the woman Miss B No. 2 is postulated, since nothing is certainly known of her in the family or in the village. Mopet oF DEATH. Vie. ACING: 1 Tee .. Died by drowning himself.* Mrs. A, nee Miss B No. 2 Death unknown. Mrs B No. 3 “- .. Shot himself. Mrs. B, nee ?—No. 4 .. Died a normal death. P, GENERATION. In this generation Miss A No. 3 is known to be a cousin of Mr. B No. 4, whom she married. He was still living in 1907, and was then 80 years of age, and had been sane all his life. This cousin marriage confirms the conclusion derived from the known *In all cases but this one, drowning was known to be by the characteristic method already described. HUMAN PEDIGREES iD existence of brothers-in-law in P,, that the originally unrelated families A and B have intermarried. Mope oF DEATH. Mrs. A, nee?—No.1 .. Is known to have been normal. Mr. A No. 2 e. .. Shot himself. Mrs. B, nee Miss A No. 3. Drowned herself. Mr. B No. 4 fe .. Was still living in 1907 and then 80 years of age. Sane throughout life. Mr. B No. 5 os .. Tried three times to shoot himself and failed. Died a normal death. In the pedigree he is marked as an Insane member. Mrs. B, nee?—No.6 .. Is known to have been normal. P GENERATION. In this generation, a second cousin marriage, resulting in a second intermarriage of the A and B families, has occurred. Some individuals of this generation, in addition to those who have actually destroyed themselves, have been temporarily placed, at one time or another, in asylums, because they became intensely melancholic and it was feared they too would destroy themselves. Move or DEArtH. Mrs. B, nee Miss A No. 1 (cousin and wife of BiNOS2) 3. .. Was temporarily in an asy- lum. In 1907 was 40 years of age and alive. 16 THE MENDEL JOURNAL Mri B No.2 ©. a Mr. B No. Mr. B No. Mr. B No. Mr. B No. aS oO -® W Miss B No. 7 Miss B No. 8 Miss BNo.9 .. Miss B No. 10 .. Miss BNo. 11 .. Mrs. B, nee ?—No. 12.. Mr. B No. 13 Miss B No. 14 .. Miss B No. 15 .. Miss B No. 16 .. Was temporarily in an asy- lum. In 1907 was 40 years of age and alive. . Unmarried. Drowned himself. . Unmarried. Drowned himself. . Poisoned himself. . Was crazy throughout manhood, but never put ‘ into an asylum. Shot himself in 1907 when 5v years of age. . Poisoned herself ‘‘ because she vowed she would never see the marriage of her brother (B No. 2) to an A No. 1, his cousin.” . Was in an asylum for a time. In 1907 she was < ce 99 = becoming “‘ queer” again. . Unmarried. Sane. . Married. Sane. . Married. Sane. Introduced by marriage ; wife of No. 13; normal. . Shot himself. . Unmarried, and was for a time in an asylum. . Sane. _ Sane and married. Re- moved with her husband and two children from their native village. HUMAN PEDIGREES 17 fr —Nonl 7 .. Husband of B No. 16, and introduced by marriage. Normal. F, GENERATION. Master B No.1 .. ; Quite children. Sane up to the Master B No.2 .. year 1907. Mr. B No. 3 .. Is now 22 years of age and in 1907 was sane. The two remaining children (Nos. 4 and 5) of this generation are quite young, and their name, sex, and age are not recorded. They have removed, with their parents, from the village, and the desire of the family is, I believe, to lose its identity. The children, none the less, constitute two of the most interesting members of the Pedigree, since they are the only members in this Pedigree both of whose parents are normal. Their future history is a matter of the greatest importance, both from the Mendelian and the medical standpoint. I am endeavouring still to . trace them. THE DEDUCTIONS FROM THE PEDIGREE.* We are not justified yet in saying dogmatically that this Pedigree is an illustration of Mendelian phenomena. But there are clearly some Mendelian indications. In the P generation all the individuals have passed the twenty-fifth year of life**, and *We follow the example admirably set by Professor Pearson, in his ‘Treasury of Human Inheritance,” in keeping our interpreta- tions apart from the facts of our Pedigrees. Those who seek for the latter can thereby obtain them without wading through the former, which at this stage of enquiry must necessarily be wholly tentative. ** The youngest must be nearly forty. B 18 THE MENDEL JOURNAL it is between this and the twentieth year that as a rule the manifestations of insanity make their appearance. Yet there are some individuals who are still sane. Others have destroyed themselves or have been confined in asylums, in order to prevent self-destruction. This fact suggests a segregation of the morbid diathesis from the normal condition. But it is only right to add that melancholia is said to be a trait of the whole present family. What such a statement means is difficult to precisely understand. Whether this melan- cholia is to be regarded as a diluted form of the insanity of the family, or whether it is a distinct morbid diathesis of itself, or whether it is merely the melancholy that is not infrequently associated with phlegmatic temperaments leading a monotonous life, or whether, in this particular case, 1t may not be merely a neurotic boding, excited by a lonely country life and a knowledge of the family history, it is impossible to say. Many families have melancholic members, but suicidal mania is not manifested by them. The facts of the pedigree, I think, justify us in believing that if a member of the family has passed the twenty-fifth or thirtieth year of life without showing any peculiarity of behaviour, he is a sane member of that family. The existence of the in- dividual, No. 6 in the P generation, who did not shoot himself until he reached fifty years of age, does not invalidate this statement, for, as is known, he was “cranky throughout his manhood.” HUMAN PEDIGREES 19 . The next feature in the Pedigree we have to consider in order to deal with it from a Mendelian standpoint is the view we ought to take of the con- dition of members Nos. 1, 2, 8, and 141in the P genera- tion. These individuals did not attempt to commit suicide, but their degree of melancholia was so great and their general behaviour such that, knowing the family history, their friends thought them safer in an asylum. In 1907 they were free citizens and were still living. In the Pedigree Chart we have indicated them, in the way their relatives and medical advisers apparently viewed them, as insane. If they are to be regarded as potentially afflicted with the family insanity, then they bring the number of insane members too high, if the case is regarded as one of simple Mendelian segregation of insanity and normality. Ii, however, they are to be regarded as merely extremely melancholic persons, but not insane, then the Mendelian expectation will be 7 -5 of those afflicted and 7-5 of those not afflicted. The result—upon this view of the nature of these members—is 6:9 respectively. (2) A Pedigree of Tuberculosis. The Family of C. Pedigree Chart 2. BY GEO. P. MUDGE. The following Family history was brought to my notice, in response to some enquiries which I made, in the early part of this year. The information was 20 THE MENDEL JOURNAL vouchsated by a member of the family, the No. 4 in the F, generation. The family is a well-to-do Irish one. It is chiefly remarkable in the apparently sporadic appearance of tuberculosis in the present or F, generation and in the manifestation of that disease, as a rule, at between 19-20 years of age. The family is one of such social position that it excludes the factors of malnutrition and unhealthy surroundings as an operative environment in the causation of the disease. THE Facts OF THE PEDIGREE. P, GENERATION. . Retired Colonel. Death unknown. . Died of senility at 80 years of age. LO. 2 2) | 29 99 » at childbirth. a P GENERATION. All normal except No. 5, who died of cancer. Nos. 4 and 6 are sisters, and No. 2 is the sister of No. 3. F, GENERATION. No. 1. Died at 18 years of age from rapid pulmonary consumption. . 2. Stillalive. Hassuffered from ulcerated throat. , 3. Died at 19 years of age from tuberculosis of stomach. ra . Quite normal. 5. Quite normal, but has gout in one finger joint. » 6. Still alive, but had tuberculous neck glands extracted at 19 years of age. 21 HUMAN PEDIGREES “ce “c UdAQS ce ‘c “ec tf ‘UOAP[IYO W1OG-[[19S XIS WOLIppe UL VIO o1OTLL, x “uossod snopnoseqny AT[e1WUe40d Ayqissod ¥v soyeoIpUT ‘soUT] SSOr0 YITA [OQUIAS OY], “S[PNPLATPUL [VULLOM 94voLpuT spoquiAs posuls oy, “Jl WOLF pop oavy AO S[soTNoteqny Jo sqyoolqns oY} Udeq GAVY OYA S[VUPLATpUL ot} eFvorpul spoquids youyq OU], ‘reok yeyy UL Ode Ia} “SUTATT [[1J8 Oso] Jo osvo UT IO “PoAT| Aoyy pvy ‘6061 avak ay} Ul poyover oAvy plNom oyefat LOY} WOM 0} S[VNPLATIPUL OY} 958 OY} OFBOIPUT WOT} OAOGB STBIOTUNU LOT]BUIS OTL ‘auole sosodand oatydiaosep 10j ore “ow “g ‘Z ‘[ S|BIOMUNT JoSIe] OL, , SISOTMOLquy, Fo a10Ip9d Ws— T1 Wd , PN Oe et ew Oe Ges 8 OR at ee (Sg Mc TE 3 age l Ql Od Go 6¢ Ee ve 9e 6L €o Go 96 86 66 Oe 0 +b+b+0+6+540 "o+6+5+5+6+6 Ps [I Moy aasbrpag 22 THE MENDEL JOURNAL No. 7. Suffers from cough, is very susceptible to cold, very slight in build, is losing flesh, is 19 years of age, but looks only 15, and has no moustache. =. i. Normal. ,» 9. Delicate: suffers from headache. 5 LO. Died at 19 years of age from tuberculosis of vertebral column. » ll. Died from tuberculosis of lungs. First manifested between the 19th and 20th year. ,, 12. Died at 13 years of age from rapid pulmonary consumption. .. 13. Quite normal. ,» 14. Normal, but suffers from adenoids. In addition to the individuals here given, there were in the left hand Family (members Nos. 2 to 7), six more children which were stillborn, and in the right hand Family (Nos. 8 to 14), there were seven stillborn children. DEDUCTIONS FROM THE PEDIGREE. One fact calls for observation. In the P genera- tion, Nos. 2 and 3 are sister and brother respectively, and Nos. 4 and 6 are sisters. Tuberculosis has ap- peared in both the branches to which Nos. 4 and 6 in part gave rise, and it has also appeared in both the branches which owe their origin to Nos. 2 and 3. Nos. 3 and 4 and 2 and 6 may therefore be tentatively regarded as D R’s, since clearly the appearance of themselves and their members in a family has resulted in the manifestation of tuberculosis in that family. HUMAN PEDIGREES 23 Ii we regard the capacity of contracting tuber- culosis as due to the absence of some factor which is present in normal individuals, then the parents Nos. 3 and 4 and 2 and 6 may be symbolised as Nn, where N-=normality, and n =absence of nor- mality =tubercular predisposition. We shall then expect in the offspring of Nos. 3 and 4, one tubercular to three normal individuals. If we bear in mind the fact that the young man, No. 7, suffers from cough, is very slight in build, is losing flesh, and looks younger than he really is, and if we further remember that he has only just reached the age, namely, nineteen, when the disease first manifests itself in this family, there can be little doubt that the prognosis in his case is a grave one. If we then regard him as potentially tubercular, that will give us 3: 3, instead of the expected ratio in this particular branch. If we make similar assumptions with regard to the constitution of the two parents Nos. 5 and 6, we find that in their offspring there are 4 normals to 3 tuberculates, instead of 5:2 to 1°8 respectively. Taking the whole F, generation together and includ- ing No. 7 among the tuberculates, we have 7 normals to 7 tuberculates, instead of 10°5 to 3°5 respectively. In regard to this discrepancy of numbers, we must remember there were thirteen children who were stillborn. Had they lived, it is possible the ratios may have been more accordant; they may have also been more discordant. We have yet to learn what is the relationship between the tubercular 24 THE MENDEL JOURNAL diathesis and the “struggle for existence ” during intra-uterine life. We have also yet to learn whether the people who are N N are more resisting in their powers than those who are Nn. For of the 15-7 persons who are expected’ to be normal in this Pedigree, only one-third will be N N in con- stitution, and the rest Nn. We are not justified in believing that both types will necessarily be identical in resisting powers. The one may resist disease under all conditions, and the other only contract it under exceptionally bad conditions, leaving it to the persons of nn constitution to fall victims under even the best conditions, foredoomed to die of tuberculosis. And, as a matter of fact, No. 6 in F, has had only a local glandular attack and is now apparently quite well. Is she anNn? If we take her and her brother, No. 7, out of the list, then we have 9:5 where 10-5:3-5 is expected. The young man No. 7 is from the modern patho- logical standpoint an interesting case, or rather will be, if tuberculosis declares itself in him. Can the hidden weakness in his constitution be stayed by an opsonic substance ? (3) A Pedigree of Human Hybrids. Segregation of European Skin Colour in a Quadroon Fraternity. BY GEO. P. MUDGE. ~ I am indebted to the courtesy of Mrs. Haig Thomas, who very kindly passed on to me some HUMAN PEDIGREES 25 information supplied by her cousin, Colonel H. de H. Haig, R.E., for the following extremely interesting case of the segregation of European skin colour in a generation of quadroons. To Colonel H. de H. Haig I am under a very great obligation for the interest he manifested in my enquiries and the great trouble he took in the endeavour to ascertain the answers to them. He very kindly sent me a photograph of a group of persons in which three of the ladies, who belong to the pedigree, appear. The facts of the pedigree are given upon the authority of Colonel H. de H. Haig, who was person- aliy acquainted with all the persons (with three excep- tions) who appear in the pedigree. I am not of course permitted to mention names, nor can I describe the geographical locality to which these events relate. It is, perhaps, permissible to say that they did not occur in the West Indies. THE Facts oF THE PEDIGREE. THe Parents =P GENERATION. The father of the family was almost certainly a yellow mulatto. His parental origin is not known for certain. His skin was light yellow brown; it ‘ was the “colour of leather called nut-brown” and “lighter than a new brown boot.” His hair and nose were quite negroid, and his lips slightly so. The mother was an Englishwoman, and a daughter of English parents of good social position. The mulatto father came to England to study medicine, and upon qualifying, he married the English 26 THE MENDEL JOURNAL lady and returned to his birthplace for the purpose of practising his profession. THE CHILDREN=F, GENERATION. There were seven children. I will describe them in order of age. No. 1 is a daughter. ‘‘ Her hair and nose are unmistakably negroid. Her skin colour is like her father’s, but lighter. Her lips are not noticeably negroid. She is very easy going and good tempered.” Nos. 2 and 3. Both were daughters. ‘ The skin colour of both is indistinguishable from that of ordinary English girls; it 1s quite white. The hair of both is typically European in form, there is no trace of the negro crimp, and in both it 1s dark brown in colour. The two girls are very beautiful and have married Eyropeans, both of them officers in the Army.” No. 4 is a son. He was not seen by Colonel H. de H. Haig, but he is known to have both skin colour and negro hair like the brother next to be described. No. 5 is a son. “ He had negro hair; the skin colour was like his father’s but lighter, though darker than that of his eldest sister.” ™m addition there are two other daughters whom my correspondent has not seen. One of these is married, but it 1s not known if she has children. THE GRAND-CHILDREN = F., GENERATION. Both the married daughters Nos. 2 and 3 have a family. One has a son and a very pretty daughter, HUMAN PEDIGREES 27 and the other has one son and three daughters. These six children have not been seen by my correspondent. THE DEDUCTIONS FROM THE PEDIGREE AND GENERAL OBSERVATIONS. There is one fact which I think needs particular notice here. In considering this pedigree, we are bound of course to face certain possibilities. Many Europeans were resident in the island. But it needs only the most casual glance at the photograph to render it at once clear that the likeness between the three sisters is very pronounced. There can be not the slightest doubt, I think, that they are sisters by the same parents. I have shown the photographs to several friends and they concur in my belief. Of course the resemblance is much greater between the two white sisters than between them and the coloured one. But even between them there is, In the form of the eyelids and the eyebrows, a strong suggestion of sisterhood. If this were a Mendelian phenomenon of the simplest order, such as Professor Pearson has imagined possible, we should expect that the offspring of such parents would be in equal numbers “ whites ” and coloured persons. There are actually 2 “ whites”: 3 coloured persons plus two of unknown type. But it is clear that this is not a simple type of Mendelian phenomenon, because the coloured children are not alike in respect of skin colour or hair characters, and they are not exactly like their father. It is clearly a more complex type of Mendelian 28 THE MENDEL JOURNAL inheritance. But whatever the nature of its com- plexity it is obviously a case of Mendelian segre- gation of European skin colour in the gametes of a coloured father. It is not necessary to consider the general problem in detail here, because it is fully considered in a later article in this Journal entitled “Skin Colour in Human Hybrids,” pp. 163. PAPERS READ TO THE MENDEL SOCIETY. THE EVOLUTION OF MAN. (An Address delivered to the Mendel Society, February, 1908.) By J. T. CUNNINGHAM, M.A., Oxon., F.Z.S. THE present paper is an attempt to consider some of the modern conceptions of Biology in relation to the human species. With regard to species in general, the Darwinian theory assumed that the differences between species were differences of adaptation, that specific characters were useful, that species were adapted to different modes of life. It has, on the other hand, been maintained by later zoologists that in the vast majority of specific characters there 1s no evidence of such utility, or of correlation with useful characters. The most eminent systematists distinguish now, as those of pre-Darwinian days did, between diagnostic characters, which are of chief systematic value, and adaptive characters, which for purposes of classifi- cation are often rather misleading than significant. The more useless a character is the more valuable it is as an indication of affinity. One modern school of 30 THE MENDEL JOURNAL evolutionists, recognising the uselessness of diagnostic characters, holds that they have not been evolved by selection, but have arisen spontaneously as muta- tions; and, with the usual tendency to carry a doctrine to extremes, they maintain that all characters are independent of utility, all arose as mutations. The American investigator, Dr. T. H. Morgan, has published a book specially devoted to this doctrine, in which he endeavours to show that adaptations do not really exist, that mutations have occurred which could only survive under special conditions of life, which in some cases the modified creatures have found, so that habits have been determined by struc- ture, not structure by habits. Thus in the short period of half a century we have had the swing of the pendulum of biological opinion from one extreme to another, from the belief that all characters were adaptive or useful to the belief that none were adap- tive. In the meantime the common-sense view has persisted that some characters were useful and some were not, and that the former were easily modified by conditions of life, the latter unaffected by such con- ditions. It must at any rate be admitted that usually in studying any group of animals we can certainly distinguish between characters which have no visible relation to the maintenance of life and others which are necessary or advantageous to that purpose ; and it is therefore possible to consider the origin of these two kinds of characters separately. The human species, in spite of the attention devoted to Anthropology, and although it is to us the EVOLUTION OF MAN 31 most familiar species, has perhaps been less studied from the zoological point of view than any other. It is also from this point of view the most difficult, partly because it is our own species and we cannot get far enough away from it to see it in true perspec- tive, partly because it has had such an exceptional history, having spread over the whole earth and become largely independent of physical conditions ; that is to say, it has attained to a great extent the power of making artificially uniform conditions which render it independent of differences of climate, geographical features, and differences of fauna and flora in different habitats. The first question to con- sider is whether man is a single species or several, and what is his relation to other species. This question, as well as most of the others which I propose to consider in this paper, has been discussed with his usual thoroughness and judgment by Darwin in his “Descent of Man,” so that I am really only trying to see whether we know any more about these prob- lems than Darwin taught us. The chief peculiarities of man, as compared with his nearest allies, the anthropoid apes, are all adaptive and useful characters, namely, the erect position, the structure of the hand and foot, and the faculty of articulate speech. Associated with the possession of language are the size and differentiation of the brain, especially of the cerebral hemispheres, and the correlated size and shape of the cranium. The reduction of the jaws, teeth, and face generally is also a characteristic feature, and adaptive to the 32 THE MENDEL JOURNAL diminution in the use made of the jaws and teeth in feeding and fighting. If man is regarded as a single species, then he affords a conspicuous instance against the doctrine that specific characters are not adap- tations, but it must be remembered that the con- tention is not that no specific characters are adaptive, but that in a vast number of cases several species are distinguished and named which live in the same district under the same conditions, and that where they live in different habitats there is no evidence that the characters correspond to differences in the mode of life. On the other hand, there is no reason why a single species should not become adapted to some peculiar mode of life, but then it would be a matter of opinion among systematists whether it should not be placed in a separate genus. | Before proceeding further with this part of the subject, it is interesting to consider the origin and nature of these adaptations. While others have been disputing whether acquired characters are ever in- herited and whether adaptations are due to the inheritance of acquired characters, Dr. Archdall Reid has made the brilliant discovery that such adaptations as those which distinguish man from the anthropoid apes are not inherited at all, but are acquired by every individual in the course of his development. Inborn or congenital characters, he says, are developed by the stimulus of nutrition alone, acquired characters are developed by the stimulus of use. Modifications acquired as a result of use and disuse are clearly never transmitted, because they never develop except in EVOLUTION OF MAN 33 response to the same stimulation as in the parent : “Plainly, then, that which is transmitted to the, infant is not the modification, but only the power of acquiring the modification under similar circumstances —a power which has undergone such an evolution in high animal organisms that in man, for instance, nearly all the developmental changes which occur between infancy and manhood are attributable to it.”* Now, while it must be admitted that it is very important to ascertain how far characters are developed entirely as the result of the constitution of the germ- plasm, and how far they require an appropriate stimulus, I think Dr. Reid attributes excessive im- portance to the latter factor. We know that a child only learns to speak by hearing speech, but we also know that monkeys and dogs do not learn to speak under the same conditions. The difference, therefore, between man and his nearest allies is not in acquire- ment, but in hereditary constitution. Indeed, Dr. Reid admits as much, but he puts the fact in other words, and here, as in much else that he has written, it seems to me that he imagines he has discovered something new when he has expressed what was known before in different and unnecessarily abundant language. He says that since “ Nature” has en- dowed animals with the power of making not all possible acquirements, but only certain fixed acquire- ments that are commonly useful to the species, there- fore species differ not only in characters which are inborn, but also in those which are acquired ; for * Principles of Heredity, 2nd Edition, page 35. c 34 THE MENDEL JOURNAL instance, the torelimbs of both ox and man grow greatly in response to use, but the lines of growth are very different. Exactly. Of course, Dr. Reid assumes that the difference in the power of making acquirements is due to natural selection. The power of growth in response to exercise resides, he says, not especially in the parts which are most used, as joints, teeth, or tongue, but in the parts in which it is most useful ; in other words, in those parts where it has been evolved, not by use, but by natural selection. It would be difficult to compress a greater number of fallacies into such few words. The chief fallacy hes in the word use. Use of a muscle means contraction, and contraction causes growth of muscle; but it is obvious that jomts do not contract, and that a joint has no size. The fact that jomts can be developed by use is proved by their actual formation occa- sionally in neglected fractures. It is also obvious that the tongue being a muscular organ is developed by exercise not merely in absolute size, but in the complexity and precision of its movements, as in the muscles of the hand; otherwise we could not learn to speak. In fact, it is precisely because the power of acquiring certain structural adaptations resides in those parts which are used for certain purposes that Lamarckians conclude that the power to acquire and the acquirement are due to the same causes ; in other words, that the hereditary or congenital factor and the acquired factor in adaptations are both due to external stimuli. The contrary view is mere assertion based on no evidence. What evidence, for EVOLUTION OF MAN 35 instance, is there that the ancestor of man possessed a variation in the power of acquiring the upright position, independently of the attempt to walk on his hind legs ? Dr. Reid supposes that this power of making acquirements is greatest among the higher animals, and little or not at all present in the lower animals and plants. He instances the frog, and expresses his belief that a tadpole enclosed in a hole or crevice, if supplied with food, would develop into a perfect frog, and that this is possibly the explanation of those cases reported in the newspapers from time to time of perfect frogs found enclosed in stone in quarries. It is unnecessary to discuss seriously this suggestion ; it will be sufficient to consider how much foundation there is for the dictum of Dr. Reid that the frog’s body gains nothing from use, and its mind almost nothing from experience. This implies that the metamorphosis is entirely due to heredity and not at all to stimulation. It has been proved, on the con- trary, that aquatic larve of Amphibians can_ be made to retain the larval state by forcing them to breathe in the water and not allowing them to breathe air, so that in this case, as in many others, the develop- ment is partly due to acquirement in Dr. Reid’s sense of the term. Dr. Reid contrasts with the supposed development of the frog the alleged fact that if the limb of an infant be locked by paralysis or by a joint disease it does not develop into an adult limb, but there is every reason to believe that the same statement would be true of the frog. 36 THE MENDEL JOURNAL We must conclude, then, that man differs from the anthropoid apes chiefly in adaptational characters, and that these characters are inborn or congenital. They are congenital in two senses; firstly, in the sense that they develop to a certain degree under what Dr. Reid calls the stimulus of nutrition, by which he means nutrition, moisture, heat, and oxygen, the essential conditions of all development and all life; secondly, that they attain their adult develop- ment from a hereditary tendency to certain modes of use and function, and from a degree of exercise which would not produce the same development in any other species. The new-born infant differs from the adult man, but it also differs from the new-born ape in all essential human characters, and that adult has acquired structural peculiarities which no ape could possibly acquire from any stimuli in its own lifetime. Obviously these are not merely specific characters, and man is not merely a species of a wider genus. Adaptational differences are characteristic among other animals of a genus, or of a family, or of larger groups. For example, among the mammals the orders are distinguished by differences of adap- tation, e.g., the Cheiroptera and Carnivora; but within a single order a family may be so separated, as in the case of the mole family. It is not easy to find a genus so distinguished. Man thus appears to have the rank of a family. The condition of the hair in man might possibly be regarded as a diagnostic character which is not adaptive; if the absence of hair on the body be explained by uselessness, still the EVOLUTION OF MAN 37 special development on the head looks like a non- adaptive feature. It is difficult, then, to regard man as merely a genus of anthropoid apes. On the other hand, we do not find that man can obviously be divided into a number of distinct species as other families of mammals can, or as even a genus can be divided. There are distinct races of man, and the question is whether these correspond to species among other animals. To discuss this question we have to consider the diagnostic characters of these races. Darwin considers them very carefully in the work I have already mentioned, and comes to two remarkable conclusions which are of chief im- portance in relation to the object of this paper— firstly, that these characters graduate into each other so that the races cannot be absolutely defined ; secondly, that they are in no sense adaptational. He says that, so far as we can judge, none of the differences between the races of man are of any direct or special service to him, nor can they be accounted for in a satisfactory manner by the direct action of the conditions of life, nor by use and disuse, nor through the principle of correlation. He then pro- ceeds to enquire whether they can. be explained by sexual selection. He concludes that this process will not explain all the differences, but that there is a residuum which must provisionally, at least, be regarded as due to spontaneous variations which have become constant and general without selection. Thus we find Darwin in this case compelled to adopt the view which in my opinion still holds good in man 38 THE MENDEL JOURNAL as in other animals, that there are two categories of characters, namely, the adaptive and the non- adaptive. The latter are of the same kind as those which are called mutations by modern biologists, while the former, in my opinion, are directly due to stimuli. Where the stimulus is functional, the modification is such as to render organs and structures more fitted for the functions ; but certain conditions may produce a direct result which has no connection with function, and which is, therefore, not in the ordinary sense useful or adaptive; for example, the absence of light stimulus causes the absence of pig- ment from the lower sides of flat fishes, but this character is neither useful nor adaptive. The real distinction between the two kinds of characters according to my views is that those of one kind are due to external stimulation, those of the other kind are independent of external causes—the latter are mutations, the former may be called modifications. Huxley’s classification of the races of mankind is a somewhat simple one. He divides them into two primary divisions, resembling perhaps genera, namely, the Ulotrichi with woolly hair, and the Leiotrichi with straight hair. The Leiotrichi he subdivides into four groups: the Australioid, the Mongoloid, the Xanthochroi or fair whites, and the Melanochroi or dark whites. The characters in which the races differ are colour of skin, hair, and eyes, shape of cranium, whether brachycephalic or dolichocephalic, character of hair, projection of jaws, shape of features, especially nose and eye-apertures. The negro race EVOLUTION OF MAN 39 is one of the most distinctly marked, its characters being black-brown skin, woolly hair, prognathous, dolichocephalic skull, thick out-turned lips, flat nose. The type is most perfectly developed in equatorial Africa, as in Guinea; to the north it has crossed with the Berbers, on the east with the Arabs ; to the south it shows reduced characters apparently without crossing. The Bushmen and Hottentots, while apparently belonging to the negro stock, are much lighter in colour, and this is some evidence that the black of the negro is originally due to the tropical sun. In the East Indian Archipelago is a type allied to the negro, but not identical, extending through the region called from its presence Melanesia, from Papua to Fiji. These have woolly hair, but the brow ridges and the nose are more prominent than in the negro ; the colour varies from black to chocolate. The Tasmanians, now extinct, are stated to have been an isolated colony of this race. The Andaman Islanders form a connecting link between the eastern blacks and the negroes, but although they have frizzled hair, which they shave off, they are brachycephalic, less prognathous than the negro, and the nose is narrower. The Australians are brown in colour with wavy black hair, dolichocephalic, having prognathous skulls with well-developed brows, and wide but not flat nose. Allied types are believed to occur in India, namely, the hill tribes descended from the primitive in- habitants, and in North-east and North Africa in the Nubians, Berbers, and the ancient Egyptians. The Mongoloid race seems to occur in its purest state in - 40 THE MENDEL JOURNAL Central and Northern Asia, and its features are : yellowish colour, long black straight hair, high cheek bones, short nose, and, especially, slanting eye aper- tures. The allied races are of immense extent and the features in many considerably modified either by variation or inter-crossing. The skulls in some of these are extremely dolichocephalic. In Asia the Mongoloid type is extended by the Japanese, Chinese, Siamese ; in Europe its invasions are repre- sented by the Turks, Finns, Lapps, and Hungarians. The Malay race seems to be a branch of the Mongo- loid, and extends over Java, Sumatra, Borneo, and can be traced to New Zealand and throughout Micronesia and Polynesia. It is supposed that the Maoris and the islanders of the Pacific, differing con- siderably from each other in characters, have arisen chiefly from crossing between a race allied to the Malays and the darker Melanesians. The inhabi- tants of the whole of America, North and South, seem to belong to one main type supposed to have been derived from the Mongolian. The uniformity of the American Indians, as compared with the diversity of types in the Old World, is one of the most striking facts in anthropology, and is most probably ex- plained by the view that America was populated from a single race, the Mongolian, from Asia, within a period so comparatively recent that no great diver- gence has been developed. Lastly, we have the white men, whose home is chiefly in Europe, and who include a great variety of subordinate types. EVOLUTION OF MAN 41 The type with blonde hair and blue eyes is found chiefly in the north, e.g., in Scandinavia and North Germany, but representatives of it are found in North Africa and in Western Asia. There can be little doubt that this is a pure type, but Huxley suggests that the Melanochroi originated from the mixture of the Xanthochroi and Australioids of India and North Africa. I can only pretend to offer a few suggestions on the characters which distinguish the races thus rapidly surveyed. Mendelians will assume that they are all simple mutations, but this does not seem to me to be a reasonable conclusion. Some of them, as in other groups of animals, are differences of degree in those adaptive characters which distinguish man from the apes. For example, prognathism and the size of the cranium and brain. The decrease in prognathism is obviously associated with the degree and duration of civilisation, so that this character in negroes and Australians is adaptive. Perhaps the same may be said of the extreme dolichocephaly associated with a sloping forehead in the same races, but we cannot say that the men with the shortest skulls are the most civilised, for some of the least civilised Mongolians are more brachycephalic than the English. In my opinion, there is good evidence that dark or black skin-colour is correlated with the light and heat of the tropical sun. It may be objected to this that the American Indians of the tropics are not black like the negroes, but they are known to be considerably darker than those of the north. At 42 THE MENDEL JOURNAL anyrate, we have the following facts: that no very dark race occurs in temperate climes either north or south, that the negroes of equatorial Africa are distinctly darker than the Bantus and Hottentots of South Africa, and that the negroes of North America have become lighter since their importation. I have sometimes thought that perhaps modifications due to stimuli might differ from mutations in not exhibiting Mendelian segregation, but the evidence so far as we have any is contradictory, for while crossing of negroes with whites always gives inter- mediates in all degrees of mixture, we have a con- stantly repeated segregation when dark whites and fair whites interbreed. Eimer mentions this as especially conspicuous in South German villages, where the inhabitants continually intermarry, and yet pure blondes and dark children occur constantly in the same family. This may be typical Mendelism, the dark complexion being dominant and the blonde recessive; but it requires further investigation.* There are, however, many race characters which seem to be evidently mutations, since there is no evidence that they are useful or due to external conditions. As examples of these, we may mention the character of the hair with regard to curling, the direction of the eye-aperture, the prominence of the nose. We have little precise evidence concerning Mendelian inheritance in these. Mr. G. P. Mudge published *Mr. C. C. Hurst has recently shown that dark eyes of any shade are dominant to blue eyes, and that the two characters segregate in Mendelian fashion.—Proc. Roy. Soc., 1908. EVOLUTION OF MAN 43 lately some data concerning the inheritance of such features in crosses between Canadian Indians and Europeans, but it seemed to me that he found all the Indian characters segregating together in one indi- vidual, and that this could only occur in a much smaller proportion of cases than he stated. His evidence would have been more convincing if he had dealt with single marked features and proved that they segregated.* In negro crosses we have no satis- factory evidence of segregation in any character. whether adaptive or otherwise. + A word or two may be devoted to the considera- tion of Darwin’s suggestion that sexual selection may account for the non-adaptive character of human races. I have shown that where the characters are confined to one sex selection cannot be the cause of this limitation. Where a character is already unisexual, however, it may vary and remain uni- sexual, as, for instance, in the human beard. The question, then, is whether selection by the female is required to account for a difference in the beard, or whether the mutation might not establish itself without selection. In deer the antlers differ in different species in size and shape, and it could scarcely be suggested that the particular size and shape in a given species was due to the fact that they * He did, however, state that segregation occurs between the blue eye, fair complexion, and light hair of a Scotsman, and the black eye, olive skin, and black hair of the Red Indian.— Nature, Nov. 7, 1907, p. 9, and Proceedings Royal Society of Medicine, Vol. IL., No. 3, p. 124. Jan. 1909. 7 This, of course, does not now hold true. See pedigree No. 3 in this Journal, page 24.—The Editor. 44 THE MENDEL JOURNAL were the best for fighting, or the most admired by the female. But sexual selection might affect characters which were not limited in inheritance ; for example, the black of negroes might be due to the preference by either or both sexes for the darkest skin, but this 1s not a probable view. In a short paper like the present I can only give a very imperfect outline of the subject, but I hope I have said enough to show that anthropology requires to be re-investigated from modern points of view. My own provisional conclusions are that man affords an example of a single species which has started a new group, which might become a genus or family. Other genera or families may have originated in this way by a single species adopting a new mode of life. The evidence does not seem to me to support the view that all human characters, adaptive and non-adaptive, can be regarded as mutations independent in their origin of habits or functional or other stimuli. The evidence seems to me to agree with the view I take of animals in general, that adaptive characters are due, not to selection, but to the effects of functional and physical stimulation, and that diagnostic characters are not adaptive and therefore not due to selection, but to blastogenic variation. Mr. Cunningham’s paper read to the Mendel Society in February 1908, was published in ‘‘Science Progress” in the following October: We are indebted to Mr. John Murray the publisher, for kind per- mission to reprint the article. - Biological Iconoclasm, Mendelian Inheritance and Human Society A PLEA FOR THE OPERATION OF A MORE VIRILE SENTIMENT IN HUMAN AFFAIRS. An Address delivered to the Mendel Socrety and to the Eugenics Education Society, in June, 1908. By GEORGE PERCIVAL MUDGE. No biologist who has lived in our community and watches its affairs, can feel that all is right in the modern sentiment that is guiding it. If he fearlessly faces the facts, and puts aside all prejudice arising out of mistaken ideas, he cannot fail to see that a con- tinuation and extension of this sentiment will lead the nation— which is Jast upon the scroll of greatness —to its destruction. Let us first consider the nature of the modern sentiment of which I speak. It is characterised by a wide range of various phases, but all of them may be expressed in one general formula, 7.e., ‘* Pre- serve and procreate the unfit citizens, and hamper and discourage the fit.” Let me illustrate the different ways in which this formula has been applied, by giving one or two examples taken from the statements and the conduct of the people who are endeavouring to apply it. When Canon Barnett says that the “weak of human society are so because of the short- comings of the strong”; when Mr. Philip Snowden 46 THE MENDEL JOURNAL says “he believes we can change human nature by education and legislation” ; when Lady Warwick and Sir John Gorst demand relief for underfed school children from the Lambeth Board of Guardians ; when certain well-intentioned but misinformed people demand that every elementary State school shall have an army of nurses to clean and bandage children’s cut fingers; when Mrs. Sidney Webb impliedly advocates that the children of workhouse paupers can be made vigorous and strong and converted into self-reliant and independent citizens if only they are nursed by State employed nurses; when Mrs. Humphry Ward believes that the children of the lower social classes are going to be made happy, well ordered, contented, and healthy citizens if the State will but provide their recreation and their playing grounds; when an Act of Parliament decrees that public servants paid out of public moneys. shall cleanse verminous persons; when certain medical men assert that the stunted weaklings whose parents live in one-roomed tenements are more stunted than those whose parents live in two-roomed tenements, and these more so than those of three-roomed tene- ments, and in consequence they conclude that our nation is undergoing physical degeneration along this road, and then further by implication recommend that the State (c.e., the fitter citizens) shall provide four-roomed tenements; when one of the leader- writers of The Times asserts that “‘ individuals, like races, are the product of the environment,” implying a direct and moulding action of the environment ; VIRILE SENTIMENT AT and when the Glasgow Board of Guardians, at the expense of the ratepayers, send the children of reprobates, drunkards, criminals, prostitutes, and general failures to be boarded out and cared for among the intelligent, virtuous, and thrifty inhabi- tants of the western coast of Scotland, in the implied belief that they will inevitably become good citizens because of the changed environment—they are expanding this remarkable formula, and are en- deavouring to teach the strange doctrine that sand can behave as granite if only both are beaten upon by the same tempestuous sea. Verily, the age of fairy tales has not yet passed away! This corybantic sentiment, so wildly solicitous of the well-being of the civically unfit, is being promulgated from every pulpit, by every newspaper, by every social reformer, by every political opportu- nist, and by every one of the great multitude of morbid and neurotic sentimentalists. It is a phase of hysteria permeating into the soul of our nation. Surely the worst thing any nation can do is to concern itself in frantic endeavours to save or pamper its weaklings and unfit. It is dangerous to ignore the clear teachings of science, that in the hereditary processes there is an irrevocable coming out at one end of the chain of generations of the characters that go in at the other; they come out not only as they went in, but numerically increased, because the individuals bearing them are multiplied in geometrical progression; they tend to spread ever wider in each successive generation. That which 48 THE MENDEL JOURNAL started in the first generation as a pair of parents may in the seventh generation become a million individuals! We are not fully aware of the silent dangers that we incur in our endeavours to save the unfit and render their lot easier. Useless as they are, and unable to contribute to national resources or progress, they become an ever-increasing source of injury and harassment to the fitter citizens, who, either voluntarily or under compulsion, contribute to their maintenance. While the artificially-endowed unfit, who are being relieved in ever-increasing degree of their responsibilities, are faithfully procreating, the artificially-disendowed fit citizens, whose respon- sibilities increase as those of the other class decrease, are limiting the number of their children. For any nation, under any circumstances, this is a dangerous condition ; but for a nation upon the walls of whose temples has been written, in the writing of con- temporary events, the warning that in the not ‘remote future she will be called upon to measure her strength with other nations upon the battlefield, it is nothing short of criminal and disastrous folly. If our nation must depart from the laisser-favre policy, and make a positive effort to determine what the qualities of its citizens shall be, the only sound line of action is to endeavour to encourage and preserve the worthier citizens, and to discourage and eliminate the less worthy ones. And _ that result can be attained without any conscious effort, for those best fitted to survive under the conditions which reign on earth will do so by purely natural VIRILE SENTIMENT 49 processes, without any artificial and compulsory interference or help. And we may be sure that these processes are benign; biological teaching is clear on that point. By these processes, throughout the whole realm of nature, the maximum amount of happiness and vigour is attained, with the minimum amount of misery or pain. Human _ interference but decreases the former, while increasing the latter. It is better, in the faith of the greater religions, to accept our destiny. I will come now to one of the principal objects of this address, 7.e., the refutation of the main as- sumption, sometimes expressed and sometimes im- plied, upon which this modern sentiment rests. It is tacitly assumed by the exponents of this senti- ment that the qualities of the individual depend upon his environment; that he is vicious because of the viciousness of his surroundings, and good because his environment is made up of good in- fluences. It is entirely wrong; there is no justi- fication for it at all in the realm of fact. The very converse is true, for in social life the environment is the product of the individual, and not wice versa. The stunted individuals are not the product of a one-roomed tenement, but the one-roomed tenement is the expression of the inherent incapacity of this race to be able to do anything better for itself. They are not undergoing physical de- generation because they are living in a one-roomed tenement—many a better man has done that— but the one-roomed condition is the natural outcome D * 50 THE MENDEL JOURNAL of their already-existing physical, moral, and in- tellectual degeneration. These degenerates are muta- tions, and breed true to their degeneracy. The qualities of an individual are determined within him; they are inherent, and depend upon the molecular architecture of the racial lines from which he has sprung. An individual is merely a piece of his racial stock; what he can do depends, not upon his environment, but upon the innate qualities which he has inherited from his stock. Let us observe that it is a potato plant, and not a dahlia, that grows out of a potato tuber; it is a dahlia, and not a potato, that grows out of a dahlia’s tuberous root-stock. A change of soil will not alter this fact. One potato may be grown among an army of dahlias, but it comes up a potato plant in spite of that. The susceptible wheat plant falls a prey to the “rust,” but the inherently immune plant growing in a field of “rusted” plants remains unattacked and healthy. From the fertilised ovum of a fish there is developed a fish, and not a bird. Transference of the bird to water, or of the fish to the skies, will not convert the one to the other. But the exponents of modern sentiment are preaching, either directly or by implication, that it will. It is a strange doctrine, and is so contrary to the common experiences of life, that it is marvellous it should ever have been believed. But I suppose there is no limit to the amount of delusion which the circumstances of a democracy, with the various appeals that are made to the uncultured and unculturable masses for their judgement, will allow to be imposed upon it. VIRILE SENTIMENT 51 Let me now proceed to describe one interesting case, to show how false is this doctrine of environ- ment, before I pass on to the second main purpose of my address, 7.e., to demonstrate the irrevocableness and inevitableness of the transmission from generation to generation of those inherent qualities that are, for all practical purposes, the sole determining cause of what an organism will be and do. In the Tyrolese valleys and elsewhere there grow two plants known as the Summer Savory and the Flax. In the valleys both these plants are green in colour. This green colour is due to a substance called chlorophyll, and one of its imherent qualities is that it can only exist as chlorophyll within a certain range of light-intensity. In a too dull light it is not formed, and in a too intense light it is destroyed. But this chlorophyll is necessary for the existence of the plant; in its absence death ensues. Now, as we move higher up a mountain the sunlight becomes more intense, because there is a less thick layer of atmosphere, with its contained water particles, to absorb a certain measure of it. Consequently at a certain height the light-intensity will be sufficient to destroy the chlorophyll and kill the plant. The hypothesis of this modern sentiment demands that these two plants, placed under the same environment, shall behave in the same way. For, assuredly, if it 1s environment which determines the qualities or behaviour of organisms, then the same environment shall produce the same response in all organisms subjected to its influence. If this be not the as- 52 THE MENDEL JOURNAL sumption underlying all efforts at social reform then what is the justification for them ? How, then, do these two plants react to a changed environment, when they are grown in the intenser sunlight that shines on the mountain slopes at a height of 2,195 metres, instead of in the shadier valley? What justification, as living organisms, do they give us for those almost frantic efforts which are designated under the collective and misleading phrase, social reform—efforts which ostensibly aim at improving the environment of the weak and unfit on the as- sumption that thereby they will be made strong and fit? No justification at all do we find in the behaviour of any living organism. Nothing, in fact, but condemnation of such efforts. For when these two plants are moved from an old environment common to them both, and are grown from seed and reared under a new and identical environment, do they both react in the same way to the same con- ditions ? Not at all! The Summer Savory changee its colour, and becomes suffused over its whole surface with a purple colouration. Careful examina- tion of the tissues of the plant shows that this new colour is developed in the most superficial tissues only, the internal tissues remaining unchanged. What is the meaning of this change of colour under the new environment of an intenser sunlight ? The purple-coloured sap which is thus called into existence possesses the power of absorbing certain of the rays of sunlight, and especially the particular ones which are most active in the destruction of the VIRILE SENTIMENT 53 plant’s green colourmg matter. The organism is thus protected from the harmful influences of an increased intensity of sunlight. Its green colouring matter—its chlorophyll—remains undamaged, and can continue to discharge those vitally-important functions that belong to it. This plant thus thrives, flowers, and seeds under the changed environment. But it is quite otherwise with the Flax. It does not develope any purple sap; it fails to respond in that way, or in any other protective way, to its changed conditions. It cannot even reach the flowering stage, for soon after it has passed the seedling stage its chlorophyll becomes destroyed by the intenser sunlight, and it perishes. Here, then, are two organisms that give us a crucial test of the validity or invalidity of this re- markable environmental doctrine. And the answer is definite and condemnatory! What an organism can and will do does not depend on the environment, but on the qualities of the organism itself. It is one of the inherent capacities of the Summer Savory to respond to this particular kind of new environment, and to adapt itself to the changed conditions. It is one of the inherent defects of the Flax that it cannot do what the Summer Savory does; and the only result of applying the environmental doctrine to it is to kill it. This is a fact which is not without its social significance. It is a fact which is not without illustration in the case of Man _ himself. Let me give the illustration, for it is highly significant and entrancingly interesting. Certain missionaries, 54 THE MENDEL JOURNAL supremely innocent of biological facts, rushed into some complex biological problems and endeavoured to civilise the Tasmanians. Acting under the im- pelling influences of mistaken ideas, but prompted by the noblest motives, they sought to change the environment of these people, and to give them what they imagined to be a better one. Their efforts were rewarded by the extinction of the Tasmanians ! In something less than fifty years the whole race of this people was eliminated, not by war, or plague, or pestilence, but by the operation of those processes that flow from the application of that crude, theoreti- cal, and sentimental conception of the power , of the environment to make good out of bad that is the dominating influence of our own social efforts. Some day we shall learn that the characters of men are relatively fixed and stable, and that they are the products of evolution under set conditions. As we find men, at any given period, in any given place, they are adapted to the particular combination of conditions under which they have evolved, and to no other. Arbitrarily change these conditions, as in the case of the Flax and the Tasmanians, and the organisms, who are thereby no longer fitted to the new conditions are destroyed. Leave them alone as Nature made them—‘ let the Tasmanians roam as they were wont, and undisturbed ”’—and they will thrive. Interfere with them, by setting up a dogma that rushes in where biological philosophy fears to tread, and the objects of our solicitude “become bewildered and dull, they lose all motives VIRILE SENTIMENT 55 for exertion, and get no new ones in their place,” and ultimately they perish. These are considerations which should be recog- nised before any endeavour is made to interfere with the conditions and natures of the citizens of our own communities. In the same way, but in lesser degree, that the mherent qualities of a Tas- manian are different from those of an Anglican Bishop, so are the inherent qualities of the lower social strata different from those of the upper strata in civilised communities. And just as the Bishop, with his higher aspirations and nobler sentiments, failed to recognise that the Tasmanian had not got them, and could not be made to have them, so certain well-intentioned, generous, impulsive, charitable, and philanthropic persons in the higher social scale fail to recognise that the aspirations and ideals of the lower classes are wholly different from their own. And just as with the Tasmanians, so in our own social efforts, disaster and destruction will follow any interference with those benign but merciless processes of Nature which allow only individuals adapted to their conditions to survive. The Tas- manians lived the life they did because their inherent desires impelled them to do so, and there was nothing in their physical environment inconsistent with them. But neither was there anything inconsistent in that physical environment with half-a-dozen other modes of living, had they chose to live them. Other ways, it is conceivable, were open to them, but they chose the particular one, and the only one, K 56 THE MENDEL JOURNAL which their inherent qualities made possible. The hfe they lived was the product of their desires—it was the outcome of their innate nature; they were not the product of their mode of life. They made their life, not their life made them. It is the same with our own social classes. The mode of life of the higher strata is the outcome of their inherent qualities in just the same way that the mode of life of the lower is the product of their born desires and capacities. Endeavour by altering the environment to compel a cultured and refined woman of the upper classes to lead the life of the slums, and we shall eliminate her race. But, clear and obvious though this is, social reformers fail to recognise the truth of the reverse process. Try to alter the environment of the lower classes, and compel them to give up their inborn habits and desires—which are the product of their line of evolution—and to lead a wholly different life, and we shall repeat the concluding chapter of Tasmanian history. The higher classes are the outcome of their evolution, and the lower of theirs. The existence of social classes is a natural fact, and the existence of different grades of social condi- tions is but the natural outcome of that fact. The social conditions are the products of the social classes, not the social classes the products of social conditions. It is in this matter that those medical men who are interested in social problems and social reformers have confounded cause and effect. And it is out of this confusion that modern social sentiment has sprung. So long as this sentiment VIRILE SENTIMENT 57 exists, and so long as it is operative in guiding and initiating social legislation, it constitutes a source of the gravest danger to the nation. It is time we awoke. It is time we turned our attention to Biology, and ¢ refused to heed any longer the “rope-dancers in the market. places.” Let us pass now from the question of environment to consider inherent qualities and the way in which, once they have come into existence, be they good or bad, they irrevocably pass on unaltered through successive generations. An organism does not mani- fest a single quality alone, but is made up of a complex combination of many qualities. Some of these qualities, be they structural or psychical, are them- selves not simple, but complex. Some of them are independent of others in the hereditary transmission, but others are correlated and always go together, in larger or smaller groups, from generation to genera- tion. An individual, like his environment, is really an aggregate of complexities bound up within each other. This fact renders all questions of social interest extremely difficult to consider, and is the best justification of the Jaisser-faire attitude. If we depart from the certain ground of Nature, and artificially touch one link in the series of com- plexities that are interlinked and make up _ the tangled chain of life, it is not possible to forecast even the immediate, much less the remote con- sequences. They are always such as we _ never expected them to be. Nevertheless, certain main trends of this chain are sufficiently clear for practical 58 THE MENDEL JOURNAL purposes, and can be seen with sufficient distinctness running through the phenomena of life. These are the facts that I would like now to bring to your attention. When a tall person marries a short person, or one of sweet temper marries one of a bad temper, there are three conceivable possibilities as to the nature of the immediate offspring. We can imagine that tallness and shortness, or sweetness and _ badness, will blend with each other in each case, and produce offspring not as tall or as sweet as one parent, nor as short or as bad-tempered as the other. Let us call this the hypothesis of “ blended inheritance.” We can, however, imagine another alternative, and conceive that some of the offspring may be like one parent and some like the other, or that, under certain circumstances, all the offspring in one generation may be like one of the parents only, those which resemble the other parent, in the particular character considered, appearing in the next or later generations. In such a case there has been no blending of the two alternative characters, for they have passed on to the next or later generations distinct from each other, each retaining its own feature of distinction. Let us call this the hypothesis of “segregated or alternative inheritance.” The third alternative is that the union of two characters in marriage shall produce an offspring showing a new quality, different from the two which by their union produced it. We may call this “diapheromorphic inheritance.” There is yet a fourth possibility, which can, however, only occur with certain characters. The bodies of VIRILE SENTIMENT 59 the offsprmg may be a mosaic of the two characters of the parents. We may call this “ mosaic, par- ticulate, or poecilodynamous inheritance.” I am not concerned now in discussing these types of inheritance, for it is to be observed they are all based upon the visible body characters. They are descriptive of the somatic features only in inheritance. My present line of enquiry will deal with somatic or body characters as incidental but still necessary features of consideration. It is with the gametic or sex-cell characters, which are, of course, invisible to us until they are manifested in the body cells, that we shall be essentially concerned. And I would like you to try and form a mental picture for yourself of an individual as a compound made up of a large number of sex-cells, which are surrounded by body cells, but which are in a sense independent of them. The body cells are the perishable cells ; they live our life for us, and then die. But the sex-cells, in a sense, are the immortal cells, for they carry on our characters from the generation in which we live to all those which succeed us. Whatever the body cells may do, there is no evidence to show that the sex-cells respond to environmental influences, in the sense that their innate qualities can be altered. Indeed, their stability is a matter of vital importance, for without it there could not exist species that, throughout long periods of time and through a wide range of geographical distribution, retain their characters unaltered. There could be no survival of the fittest, if the qualities which made a race the fittest were not stable and, therefore, persistent. 60 THE MENDEL JOURNAL The stability of the sex-cell is a fact that I would like you to grasp, because it is one of very great importance in considering social problems. What we have always to bear in mind is not so mucha question of what the individual himself may be, but what is the nature of the characters which his sex- cells are carrying and transmitting to successive generations. And there is another fact of equal importance which we should endeavour to fully understand now, and this is that the body or visible characters of an individual are not necessarily a reliable indication of the characters which his sex-cells may be carrying. Three grey-coated animals, externally similar, may be carrying in their gametes (sex-cells) very different qualities, so far as colour is concerned. One may carry nothing but greyness, another greyness and albinism, and the third may carry greyness, blackness, and albinism. The offspring of these three apparently identical individuals would be very different. If these processes are at work in human life—and I shall presently show you evidence indicating that they are—clearly it is a matter of the widest significance, and one which all social reformers have overlooked. If a youth from the lower classes manifests a few superficial accomplish- ments—can, for instance, pass an elementary scholar- ship examination by a process of cram—but is carrying the civic qualities of his class, is it worth the while of a State to spend the money of better citizens upon that youth? Is it worth while to spend money in order that we may produce in an individual certain - VIRILE SENTIMENT 61 artificial accomplishments which can always be had for nothing in the class immediately above that to which he belongs? In this next higher grade we not only obtain the artificial accomplishments, but in addition better innate civic qualities. What we are really doing by such methods is paying for an inferior stock when we have already the better one for nothing. We pay for a few putative accomplish- ments in an individual whose sex-cells are going to give us an inferior civic stock! That is one con- sideration which this new fact of heredity bids us heed. We are going to deal, then, with the sex-cells. When the male sex-cell unites with the female sex- cell there is produced the first body cell, by the repeated divisions and growth of which a new in- dividual arises. What this new individual will. be, and what he will transmit to the next generation, depends not upon his environment, but upon what the two sex-cells, male and female, of his parents brought in. A sex-cell which carries leaden social instincts is not going to develope into a golden individual because he has been given a golden en- vironment. From it a person of leaden instincts only will develope. Let us appeal, then, to accurately-conducted experiments, and see how far we can ascertain some reliable facts as to what the sex-cells are really doing. When a sex-cell of a pure black rabbit is united with one of an albino rabbit, to what sort of offspring will it give rise? Without exception, whenever this cross has been made, black rabbits indistinguishable 62 THE MENDEL JOURNAL from the pure black parent have been the result. The whiteness of the albino parent is temporarily lost to view; but it is not destroyed or swamped. It exists, and under the appointed conditions which Nature has determined and we have now learned it will reappear, pure and unaltered. Its association with blackness, in the processes of heredity, has not altered it. Environment, even of this intimate kind, has not changed it. For, if we cross the sex-cells of the brothers and sisters of this first generation, we find that two kinds of rabbits are produced, «e., black and albinoes. Blackness and albinism have separated out again, perfectly distinct. There has been no blending of these characters, but segregation of them. We know that each member of the offspring in the first generation which results from uniting the sex-cells of a pure black rabbit with those of an albino must have been compound individuals, because each must have contamed both blackness and albinism. They are, with respect to these two characters, impure individuals. We may call them hybrids, or heterozygotes*. But the feature of interest is that although they are impure, since they carry blackness and whiteness, they are indistin- guishable from the pure black parent which carried blackness only. Something must have happened to the albinism, since it does not manifest itself in these hybrids. We are not sure what does happen to it, but it does not, for our present purpose, very much matter. We have got the fact that one of the *A term applied by Prof. Bateson. VIRILE SENTIMENT 63 two alternative characters exhibits itself in the hybrid, and the other does not. We may speak of the character which thus shows itself to the exclusion of the other as the Dominant one, and the one which temporarily disappears as the Recessive character. When these hybrids are mated with each other, or are mated back with either of their two aprents, or with individuals like their parents, blackness and albinism separate out again in this second generation. But we know from experiment that all the black individuals of the second generation are not alike. This is a very important fact, and its social bearing we have already glanced at (pp. 60 and 61). Some are pure black individuals, and others are hybrids ; the one carries blackness only, and the other both blackness and albinism. And there is a certain proportion in which these types appear when hybrids are mated inter se. It is one pure black, two impure blacks, and one albino. It the pure blacks be bred with each other, only pure black individuals are produced in the offspring. The dominant character breeds true. If the albinoes be mated with each other, only albino offspring are produced. The recessive character also breeds true. But if the impure blacks be bred together, then the offspring again consists in this third generation of different individuals in the proportion of one pure black, two impure blacks, and one albino. The hybrids do not breed true. Now a moment’s thought will show us that we cannot explain these facts on any assumption of G4 THE MENDEL JOURNAL the blending of the characters carried by the sex-cells of the two parents. Had blending occurred, black- ness as such, and whiteness as such, would have for ever disappeared. When blackness is crossed with whiteness, if blending takes place, we should expect to find a diluted blackness. And the diluted blackness of the first generation, when bred with diluted blackness, should on this hypothesis give no other individuals but those of diluted blackness; and it should do this for generation after generation. When crossed with albinism, a dilute black should give a diluter black, but not blackness as black as the original, and albinism as pure as the first albino introduced. It is clear, then, that the blended hypothesis of inheritance fails completely in this case. And it similarly fails in a large number of other cases, comprising a wide range of characters. Will the hypothesis of segregation give us an ex- planation of the ascertained facts? Assuredly it does. Let us proceed to see how. The doctrine of segregation assumes as a fundamental proposition that when two sex-cells unite, one carrying black- ness and the other albinism, these two qualities do not blend or fuse in the single cell which results, but remain distinct; and that at the ultimate cell-divisions by which the sex-cells of the new individual will separate out from its body cells, these two qualities will be found distinct from each other, and carried in different sex-cells. It the in- dividual is carrying only blackness, then, of course, all its sex-cells will carry blackness. And, similarly, VIRILE SENTIMENT 65 if it is carrying only albinism, then all its sex-cells ean carry albinism alone. But if it carries both blackness and albinism, then one-half of its sex-cells will carry one character, and the other half will carry the other. Now it is clear that so long as a pure black individual is crossed with another pure black, sex-cells carrying blackness only can meet. Hence, only a black offspring can result. And in the same way, so long as an albino is crossed with an albino, there is only one kind of sex-cell which can meet, a.e., an albinism-carrying one with an albinism- carrying one. But it is otherwise when an impure or hybrid black is mated with a similar individual. Here the two parents are both carrying two kinds of sex-cells, z.e., those transmitting blackness and those transmitting albinism. Consequently, at sexual conjugation, a male sex-cell which carries blackness has an equal chance of meeting an egg carrying either blackness or albinism. And the same is true of the male sex-cell which carries albinism. If we agree to let A stand for albinism, and B for blackness, then BA will represent a hybrid individual. This individual will be black in colour, although it is carrying albinism, because we have seen that black- ness is dominant and albinism is recessive. We may, therefore, graphically represent the possibilities in this way :— | PieAe < Bb Parents. | |BA+BA+BA | Offspring. 66 THE MENDEL JOURNAL If No. 1 BA, the sister, is mated with No. 2 BA the brother, then the B sex-cells of the first may meet the B and A sex-cells of the second, and give us BB and BA respectively. Similarly, the A sex-cells of No. 1 may meet the B and A sex-cells of No. 2 and give us BA and AA respectively. Thus we shall obtam 1 BB +22BA+ 1AA. Thatasian the average in every four members of the offspring _of two BA parents, we shall expect 1 pure black, 1 pure albino, and 2 impure or hybrid black imdi- viduals. And this is what experiment does give us. We have spoken so far of the segregation of these two characters. But there is another feature which we should note, and it is this: Segregation of charac- ters alone would not be sufficient to explain the experimental results. It is possible to have segrega- . tion and yet have both characters carried in the same sex-cell; they can he side by side with each other in the cell. But it is a fundamental part of our proposition that any one sex-cell can carry one only of the two alternative characters. To this . conception of the structure of the sex-cell we apply the term “gametic purity.” Necessarily, gametic purity implies segregation. But segregation does not necessarily imply gametic purity. We may now widen the description of our principle of heredity, and call it the theory of segregation and gametic purity. We have arrived, then, at this position: The sex-cells are the carriers of the characters of the race, from one generation to another. Whenever two alternative characters, such as tallness and VIRILE SENTIMENT 67 dwarfiness, colour and albinism, short-haired coat and long-haired coat, hairy epidermis and smooth epidermis, blue eyes and brown eyes, and a large number of other alternatives are present in a race, each of the sex-cells can carry one only of these two characters, the other character being carried by another cell. And the distribution of these two characters among the whole mass of the sex-cells of any individual is such that one-half of the total number of them will carry the one character, and the other half will carry the other character. And the outcome of this structure and arrangement of the sex-cells is that when two hybrids are mated, their offspring will consist of a mixture of individuals, some showing the dominant character and others the recessive. Those which exhibit the dominant character are not alike, some of them being pure dominants and the others impure or hybrid dominants. The proportions in which these types appear are one pure dominant, two hybrid dominants, and one recessive. I would like now to ask your consideration of a type of mating which is very important from the social standpoint, and from the point of view of the student of human heredity. It is the mating where one of the two parents is a hybrid and the other a recessive. In our symbolic representation we may state it this way: DRx RR. The result of such a mating is to produce only two kinds of off- spring, being identical in regard to the two _par- ticular characters under consideration with the 68 THE MENDEL JOURNAL two parents. That is, the offspring will consist, in the long run, of an equal number of individuals who are hybrids, or D R#’s, and of pure individuals, or R R's, who are carrying only the recessive char- acter. The hybrid individual will carry both the dominant and _ recessive character, but will ex- hibit in his structural features or psychical con- duct only the dominant one. That is, he is some- thing different from what he appears to be. His influence on the race, in respect of his hereditary transmission, may be quite the opposite to that expected, if the expectation be based on his present apparent condition. This is a fact which social reformers and all others who aspire to interfere with Nature’s processes would do well to clearly realise. It is easy enough to heedlessly interfere with the workings of Nature, and to misunderstand her, but it 1s impossible to escape the consequences of so doing. If this conception of segregation and gametic purity be true of human life, it is easy to see how important a bearing it has upon the problems with which Genetics* and Eugenics are concerned. Ii we, as students of these branches of biological science, desire to see only our individuals of good civic qualities mated, and if we find upon further study that a certain proportion of our apparently good citizens are carrying in a recessive condition the antithetic bad qualities, which may become mani- fested in the next generation, clearly our line of * For a definition of Genetics and Eugenics, see Appendix, p p. 110 and 112, VIRILE SENTIMENT 69 action or of social education must be modified. There is a pitfall lying across our path, and we must be careful to see it, in order that we may avoid the consequences of falling into it. Before I pass on to consider the other main purpose of this paper, 7.e., the question whether the evidence justifies us in believing that the processes of segrega- tion and gametic purity are operating in human heredity, I would like to make the task a little easier by tabulating all the possible types of mating which can occur. We are assuming for the present that human beings are composed of a number of alternative or unit characters. These are disposed in pairs, and each unit character in a pair is capable of re- placing the alternative one in the processes of here- dity. One of these unit characters we call dominant, and the other recessive. This assumption being tentatively accepted, we may classify human matings, so far as particular pairs of characters are concerned, in the following way, and from such matings offspring of the nature indicated will be expected :— Parent x Parent. ‘Nature and proportion of expected Offspring. fee x ey = All i. UD kh f= ADF and all-visibly D. Dx Diy = D D+ DF =-all visibly D. Dip DR = AD D412 DE (visibly D) +14 A. De x fe 1 DR (visibly D)+1 RR. EE 1) = VAM DD, Now we are in a position to consider what is, next to the question of man’s origin and destiny, one of the most intensely interesting of human questions: Do the alternative characters which make up human beings, and which decide what we l | 70 THE MENDEL JOURNAL shall be and do, separate from each other during the processes and structural changes by which the sex-cells are produced, so that, individually, we represent the product of a sum of paired characters handed down to us by our parents? And what we shall be, therefore, will depend not upon what our ancestors as an aggregate were, but upon the gametic nature and structure of our parents? And since our parents can only carry between them a single pair of alternative characters of the same class, while all our ancestors between them may have carried many such pairs, it is clear that we individually cannot represent in any particular character more than two of our ancestors, while in some cases we may represent only one and carry in a recessive condition the alternative unit character of one other? Or, are we the product of the blending of all the ancestral alterna- tive qualities that have been brought into our line by all our ancestors ? So far as our evidence is definite at all, the answer for the latter question is negative, and affirmative for the former. I am far from asserting that our evidence is complete, or that it is of such a nature that the blended hypothesis of inheritance is yet altogether excluded. But I am prepared to maintain that, making due allowances for the circumstances under which the human pedigrees have been obtaimed, and giving proper consideration to the probable complexity of many human characters supposed to be simple, the evidence is very clearly suggestive—and in some cases it amounts to a scientific demonstration—that gametic VIRILE SENTIMENT 71 purity and segregation are the two processes which determine human heredity, alike in the transmission of normal and abnormal characters. Let us pass on, therefore, to consider some of the evidence for segregation and gametic purity in man. I propose first to deal with the transmission of a normal human character, 7.e., eye-colour. Until Mr. Hurst, during the years 1905-1907, examined a number of families, separately and in detail, recording each individual eye-colour, it cannot be said that our knowledge of this matter was anything like definite or satisfactory. Previous records largely depended upon the observations of different ob- servers, and upon a more or less popular and in- definite classification of eye-colours. And, moreover, the mathematical methods that were employed to deal with the ascertained data were of such a nature that the truth was rather masked than elucidated by them. As a matter of subsequent knowledge, they actually did miss the truth. Mr. Hurst made a personal examination of the eye-colours of one hundred and thirty-nine pairs of parents, and of six hundred and eighty-three of their offspring. He found that he was enabled to classify all eye- colours into two classes, which he called the ‘“‘ Sim- plex”? and the ‘“ Duplex.” These two types of eyes are quite distinct. The simplex type includes all the pure blue and pure grey* eyes, 7.e., blue and * Pure grey eyes are simply blue eyes in which the iris tissues are a little more opaque, and so cause the blue to appear grey. Eyes which are blue in childhood may become grey in later life, owing to an increase of this opacity. 72 THE MENDEL JOURNAL grey eyes which contain no visible trace of yellow or brown pigment. The duplex type includes the black, brown, hazel, green, and impure grey and impure blue eyes, «.e., grey and blue with a little visible yellow and brown pigments present. He found that the simplex type behaves as a recessive to the duplex type; so that, if one parent had pure duplex eyes and the other had simplex eyes, all the offspring would show duplex eyes. If both parents were of the simplex type, then all the children had simplex eyes. Since the duplex eyes are a dominant type, individuals who possess them may be either carrying duplex only, or they may carry as well the simplex type as a recessive. When both parents have duplex eyes, there are therefore three possible combinations which have to be considered. We may state them symbolically thus :— DDD; DD sR Dy ken Dl) ae That is, we may have a pure duplex married to a pure duplex, or a pure duplex married to a hybrid duplex, or both parents may be of the hybrid duplex type. If we turn back to the tabulation on page 69, we can see the nature and proportion of the expected offspring in these three kinds of marriages, on the assumption that the processes of gametic purity and segregation are operating in Man. Let us consider Mr. Hurst’s results. When both parents were simplex all the offspring, one hundred and one in number, belonging to twenty pairs of parents, were simplex also. That is, the recessive character breeds true, in spite of the fact that some VIRILE SENTIMENT 73 of the grandparents had duplex eyes. This is quite in accordance with the theory of gametic purity and segregation, and it is altogether discordant with any other theory of inheritance, whether bio- metrical (mathematical) or otherwise. Turning now to the results when both parents are duplex, we have just seen that three combinations are possible. Of these, the two combinations DDx DD and DD x DR will give the same visible result, 2.e., all the offspring will show the duplex type of eye. But the third combination, ve, DR x DR, will give a mixed offspring, in the proportion of three individuals having duplex eyes (=1 DD +2 DR) to one having the simplex eyes. Examining Mr. Hurst’s results, we find that our expectations are fulfilled, for of the fifty pairs of parents which he recorded, both of whom had duplex eyes, thirty-seven of them had a total offspring, one hundred and ninety-five in number, all with duplex eyes. They, therefore, illustrate the two first com- binations. The remaining thirteen parents had a mixed offspring, of a total number of sixty-three individuals. Forty-five of these showed the duplex eyes, and eighteen of them the simplex eyes. In nearest round numbers the expectation is forty-seven and sixteen respectively. That is, Mendelian pre- diction and result are practically identical. The next possible combination will be between parents having duplex eyes and those with simplex eyes. But the duplex parent may be one of two things—either a DD or a DR. So that from 74 THE MENDEL JOURNAL the standpoint of the type of ofisprmg which we may expect, we have two real combinations within this one visible one, 2.2, DD x RR and DR x RR. In the first case all the ofispring are expected to have duplex eyes, and in the latter one-half is expected to show the duplex eyes and the other half simplex. What are the actual results ? There were seventeen pairs of parents, of the nature of Duplex = Simplex, with a total of sixty-six offspring all showing the duplex type of eye. And there were fifty-two other pairs of parents of the same visible nature who had | a total ofispring of two hundred and fifty-eight, one hundred and thirty-seven of whom had the simplex eye and one hundred and twenty-one the duplex eye. The expectation is one hundred and twenty-nine of both types. There is thus a very near approximation of result to prediction. Mr. Hurst’s work shows two things quite clearly. It demonstrates the absolute segregation of these two types of eyes in human inheritance, and compels us to accept the conception of gametic purity as the only valid explanation of it. It is quite evident that the alternative hypothesis of blended inheritance does not explain a single feature of the phenomena. For if it did, why do blue-eyed (simplex type) offspring come from two brown-eyed (duplex) parents? Or, why do they come from parents one of whom has brown eyes and the other blue ones? Why do some brown-eyed pairs of parents give nothing but dark- eyed offspring, and other pairs give a mixed offspring of dark-eyed and blue-eyed individuals ? The con- VIRILE SENTIMENT * 75 ception of blending does not explain it, while that of gametic purity and segregation enables us to form a mental picture of the main line of the processes concerned. The same segregation of eye-colour occurs when distinct races of mankind are crossed. Among my collection of human pedigrees I have several of an European (usually a fair-complexioned, light-haired, blue-eyed Scotchman) married to a Red Indian woman, and of their descendants. When the half-breeds from such a marriage marry an Kuropean of a certain gametic structure, the fair complexion, light hair, and the blue eyes segregate out again among their children. I pass next to consider another character which we cannot regard as pathological, but yet is not normal. We may speak of it as abnormal. I allude to the quality of albinism. It is a condition in which there is a complete absence of pigmentation in the body.* It is known not only among the less deeply pigmented races of mankind, but also among the intensely pigmented races, such as negroes. Mr. Farabee has examined an interesting case of the hereditary transmission of albinism among negroes. An albino negro married a normally-pigmented negress. There were three sons, all normally pig- mented. All three married pigmented negresses, and two of them had only the normally-pigmented off- spring. The other son married twice, and in each *The pigments of the blood, bile, and of muscle are, of course, excepted. It may perhaps be better, for practical purposes, to define an albino as an individual in whom there is absence of visible pig- mentation. 76 THE MENDEL JOURNAL case married a pigmented negress. But by his first wife he had one albino and five pigmented children, and by his second wife three albino and six pigmented children. The complete disappearance of the al- binism in the first generation, and its reappearance in the second, shows that albinism in man, as in lower forms, 1s a recessive character to pigmentation. The results of these marriages are perfectly unintelli- gible on the conception that blending of characters occurs. For how, by any process of blending, can four albino children be produced out of a marriage of a normally-pigmented negro and a negress? The blending of black with black does not produce white ! And how can we explain the mixed nature of the offsprmg—some individuals being albinoes and others pigmented—on the blending conception. A blend is expected to produce an uniform result. But the whole of the facts are explicable on the hypothesis of segregation and gametic purity. The albino grand- father married a normal negress. All three of their sons will be, therefore, hybrids of the nature D Rk, where D stands for the dominant character of pig- mentation, and # the recessive character of albinism. So long as these sons marry only normal negresses (i.e., D D’s), a pigmented offspring only is expected. That appears to have been the case with two of the sons. But if the third D # son shall marry a negress who is also DR, then we expect in the offspring albinoes and pigmented individuals in the proportions of 1: 3. This appears to be the kind of marriage that the third son contracted with both wives. VIRILE SENTIMENT 17 And we note that the number of albinoes and pig- mented individuals, 7.e., 4 and 11 respectively, is as near as it can be to the expected 4: 12 ratio. This case, then, must be accepted as demonstrating the existence of the processes of segregation and gametic purity in the hereditary transmission of at least one other human character. Let us consider another case of albinism, but this time among Europeans. In a Swiss village there once lived an albino, Josephine Chassot. She never married, and for three generations back her ancestors were apparently normal people. But in her own generation she had ten cousins in the second degree.* Of these, three were albinoes, and the remaining seven were all apparently normal. All the normal ones appeared in one family, and all the three albinoes in another. In this latter family it is known that there were normal members, but the number is unknown. One of the normal cousins of Josephine married a normal woman, and had twelve children, eight beg normal and four albinoes. Now this case, apparently remarkable because of the sporadic appearance of the albinoes, is entirely explicable on the Mendelian explanation of segregation and gametic purity. The families concerned lived in two Swiss villages half a league from each other, and we may suppose that at the time (¢.e., sixty years ago) travel- ling was difficult, and intercourse with more distant villages was more or less a rare event. Under such * That is to say, they were the children of two pairs of parents, one of whom in each pair was a cousin of one of her parents. 78 THE MENDEL JOURNAL conditions, consanguineous marriages are apt to be common. This we see in a marked degree in the Scotch Highlands, where the inhabitants of many villages are nearly all cousins, in nearer or remote relationship. We know that albinism is a recessive character in animals, and we have just seen in the negro marriage that it is so alsoim man. Individuals, apparently quite normal externally, may, therefore, carry albinism as a recessive character. So long as such individuals marry a quite normal person (one not carrying albinism recessive), albinism will never appear among the offsprmg. But if they marry another person similar to themselves—that is, one carrying albinism recessive—then albino children, as well as normal ones, will be born to them. So that it may happen that in a long line of ancestry, apparently normal all along, albinoes may quite suddenly arise. That appears to be the explanation of the case of Josephine Chassot. And the same explanation suffices for the familar fact that albinoes are very olten the offspring of a cousin marriage. In these villages, where we may assume consanguineous or endogamous marriages* were common, there must have existed a certain proportion of the villagers who were carrying albinism as a recessive character. When two such married—and in random selections this must now and then occur—albinoes are expected. This, then, being the Mendelian expectation with = In this particular pedigree it is known that two normal brothers married two normal sisters, all of the same village, and albinoes appeared in the descendants of both. VIRILE SENTIMENT 79 regard to a qualitative result, how far is the ex- pectation similarly corroborated by a quantitative result ? The Mendelian expectation with regard to the proportion of albinoes and normal individuals is as 1: 3. The total number of offspring concerned for purposes of calculation is twenty-four plus a few normals, the number of which is not recorded. The expectation, therefore, in round numbers is_ six albinoes and eighteen normals. The actual number is eight albinoes and sixteen normals. There is thus what the prediction requires, a large excess of normals over albinoes. And when we bear in mind that a few more normals, who are known to have existed, must be added to the actual result, we see how well result and prediction meet each other. Dr. A. M. Gossage has quite recently made a study of some previously recorded cases of hereditary transmission of pathological characters in man, and he has produced a number of interesting facts. On the whole the cases which he has studied and re- published confirm the Mendelian generalisation, though there are some exceptions which are discordant with it. These latter may, however, receive an adequate explanation when we know more of the particular diseases with which they are connected, or when we have been able to investigate a larger number of similar cases. But it seems quite clear from his survey of cases that segregation of alter- native characters, such as some pathological trait and normality, does occur. The crucial test that such segregation has actually occurred is to be found in 80 THE MENDEL JOURNAL the persistency with which a recessive character breeds true when once it has been segregated out from association with a dominant character. It is true that, with the exception of one remarkable case, which we shall consider later, we have no very long continuous pedigree of an extracted recessive charac- ter. But we have a number of short pedigrees, and when all their indications point in one direction, and almost invariably show that an extracted reces- sive character breeds true when the individual manifesting it is mated with another also showing the recessive character, we are justified in accepting this as clear evidence of segregation. And the justification is not lessened because a few exceptions occur. These may receive an explanation when we know what it is that determines which of two alter- native characters shall be the dominant one, when we have tested the validity of past records by a more searching examination of future cases, and when pathologists and physiological chemists can tell us more of the nature of the factors which go to the making up of these pathological characters. Many abnormal characters are doubtless more or less complex in their nature. To treat such abnormalities as simple units is but to disguise the truth. When we know the nature of their complexities, the general trend of the whole evidence leads us to believe that they will fall into line with the general body of facts of Mendelian inheritance. Some of the characters studied by Dr. Gossage are of suggestive interest. In a certain family VIRILE SENTIMENT 81 some of the individuals possessed hair presumably of the EKuropean type, but others had a “ tightly- curled, short, woolly hair of the negro type.” The facial features of these latter individuals were not in the least negroid. A tradition in the family attributes the woolly hair to a Mexican ancestor several genera- tions back. We may assume it as probable that this ancestor was of the negroid race. If this is so we have a clean segregation of European facial characters from negroid ones, for none of the present family, thirty-six im number, manifest the least negroid trait. But in the random segregation of the various negroid and Kuropean characters that oc- curred in the sex-cells of this Mexican ancestry, negroid hair became associated with European facial characters. And apparently, the European alter- native characters* being recessive, once the Kuropean facial characteristics had been extracted they would henceforth breed true. Hence in none of the offspring that have descended from a Mexican ancestor who had European features but negroid hair, do we expect any negroid facial characters to reappear. But the negroid hair being a dominant character, this Mexican individual who bore it may also bear in his sex-cells the European type of hair as a recessive character. We should then expect to see both the negroid and Huropean hair separate out in the succeeding generations, and be borne by separate individuals. And the curliness and woolliness must, on the whole, be as curly and woolly in one individual as another, And such appears to be the case in ‘ 82 THE MENDEL JOURNAL this family. There have been three marriages of individuals who had woolly hair ( =D R) with persons having European hair (=R R&R). The total number of individuals concerned is thirty-six. Of these, eighteen had negroid hair and eighteen had ordinary hair. The segregation is complete, and the pro- | portions are in exact accordance with Mendelian expectation. Another case in which a peculiarity of the hair is the character considered is one where a congenital tuft of white hair over the brow was hereditarily transmitted through six generations. Only a single member, the father, in each of the first three genera- tions is known. And each of these possessed the white tuft. The father of the third generation married a normal person, and they had four children, two with and two without the tuft. The white tuft is a dominant character and the absence of it a recessive one. One of the normal offspring in time married a normal husband, and their children, three in number, were all normals. The two children with the tuft also married, and they had a mixed offspring, some of the individuals having the tuft and others not. Three individuals of this generation without the tuft married normal persons, and in their total offsprmg of eight members there was not a single one with the tuft. Thus in the total of eleven children born of four pairs of parents, one in each pair being an extracted recessive and the other a normal person, the dominant character did not appear. That is, the Mendelian expectation that the VIRILE SENTIMENT 83 recessive character would breed true has been fulfilled in fact. With regard to the proportions in which the two kinds of offspring derived from parents, one of which manifests the tuft (=D FR) and the other does not (=R R), the actual proportion is nearly identical with that required by expectation. There are thirty-two individuals in the offspring, and the expectation is sixteen of each type. There are, actually, seventeen with the tuft of hair and fifteen without. I have taken these two cases from Dr. Gossage’s paper because they are of a nature which the layman can understand. They do not deal with patho- logical characters, but rather with what we may term peculiarities. But all the rest of the cases with which he deals are of a pathological nature. The phenomena of segregation and gametic purity are clearly manifested, but, as already stated, there are a few doubtful exceptions. Those who desire to study these cases will find them described in the Quarterly Journal of Medicine, Vol. 1, No. 3. I would lke now to ask you to consider two extremely important and interesting cases. The first one has reference to an abnormal condition of the fingers and toes. The peculiarities connected with them are correlated with others, but these we need not consider. The case has been worked out, with very great care and labour, by Dr. Drinkwater. The peculiarity is technically called ‘“ Brachy- dactyly’; more popularly we may speak of it as “ short-fingerness ” or ‘‘thumb-fingerness.” The 84 THE MENDEL JOURNAL fingers of the hand are apparently, like the thumb, made up of two joints, instead of three asin the normal hand. It is the middle one of the three jomts which has to all appearance disappeared. There is some reason to believe that it has not actually disappeared, but that it is vestigal in size and has become fused on to the last joint. However, for practical purposes we may speak of the peculiarity as a two-jomted condition of the fingers. The history of the family in which this peculiar condition is present is known for seven generations. The first three generations are incomplete, but the remaining four are complete and comprise a fairly numerous collection of individuals. The whole seven generations include one hundred and seventy- four persons. We may divide the marriages into two classes, 7.e., those in which an abnormal person married a normal one, and those in which an extracted normal individual derived from this family married a normal one from outside of it. The first type of marriage will be represented as D Rx R R, and the latter type by RRxRR. The brachydactylous condition is shown by the observations on’ this family to be a dominant one. Now, what is the Mendelian expectation as to the offspring from these two types of marriage ? In the first type two classes of individuals, 7.e., short-fingered and normal-fingered, are expected in the offspring. And it is further expected that these two classes will be present in approximately equal numbers. Once more an appeal to the facts of human pedigrees confirms Mendelian VIRILE SENTIMENT 85 prediction. Both classes of individuals are present, and in very nearly equal numbers. There are eighty individuals from this type of marriage, and therefore it is expected that there will be forty each of the abnormal and normal individuals. There are respec- tively forty-two and thirty-eight. With regard to the second type of marriage, ue, RRxRKR, the expectation is that all the offspring will be normal. There are eighty individuals,* all normal. The family which we have just considered is an English one. But a case of the same nature, dealing with an American family, was earlier published by Mr. Farabee. In this case the same principles are manifested. The extracted recessive (normal) charac- ter breeds true, and the offspring from a marriage of a parent with “ short fingers ” with one of normal fingers is composed of the two types of individuals, and in approximately equal numbers, 7.e., thirty-six of abnormal individuals to thirty-three of normal ones, the expectation being thirty-four and a half of each. These two cases of human brachydactyly, then, are clearly Mendelian in nature. Segregation and gametic purity explain the phenomena presented by them, while the blending hypothesis and the biometrical > “Law of Ancestral Inheritance” completely fail. I pass now to the second and perhaps one of the most important cases of a human pedigree. We owe its compilation in part to a French Army surgeon, the late M. Florent Cunier, and in part to an English * In addition to these there are a few individuals known to have existed, but the sex and type is now not known. 86 THE MENDEL JOURNAL ophthalmic surgeon, Mr. Nettleship. It is one of the largest known human pedigrees of its kind, comprising somewhere near a total of two thousand individuals, and it extends back through ten generations to the year 1637, when the individual Jean Nougaret, with whom it commences, was born. The peculiar here- ditary character which affected one hundred and thirty-five persons of this family, and which mani- fested itself in each generation, is one which is known as congenital stationary night-blindness. The real nature of the disease is quite well confirmed, for Mr. Nettleship, with Professor Truc and M. Capion, have been able to examine fifteen night-blind subjects, belonging to the ninth, eighth, seventh, and sixth generations. The affected persons can see as well during the daytime as normal people. But at night time they can see only by candle light and by very bright moonlight. On moonless nights, or in very dark places such as cellars, they are blind. The affection is present from birth, and mothers are able to determine whether their babies are night-blind or normal by the apathy or interest manifested by them when objects likely to attract their attention are held up at night time. In this case of congenital night- blindness it is shown by the facts of the pedigree that the abnormal condition is the dominant one, and the normal one is recessive. The Mendelian expectation therefore is that when two extracted normal persons marry, or when an extracted normal person marries a normal individual, that all the off- spring shall be normal. Is the Mendelian expectation VIRILE SENTIMENT 87 , tulfilled in this case, too? The answeris Yes. When- ever normal parents have married—and in this family inter-marriages between extracted normals have occurred—the offspring is always normal. There is altogether an offspring of one thousand six hundred and sixty-three normal individuals derived from the marriages of normal parents, and not a single abnormal person has occurred. As far as human investigation can be concerned, nothing can be clearer than the meaning of such a record. The segregation of the normal character, once it has occurred, is complete, notwithstanding that inter- marriages between extracted recessive individuals have occurred. And there are cases within this family where this pure breeding has extended con- tinuously for seven generations. There is nothing to suggest that the recessive character will not con- tinue so to breed for indefinite generations. When we examine the records of the offspring derived from the marriage of abnormal with narmal persons (that is, marriages of the type DR«x Rf R), we find the Mendelian expectation fulfilled in that the expected two types, 7.e., the night-blind person and the normal person, are present. But, however, the expected equality of the two is widely departed from. There are three hundred and sixty-four persons in the disease-bearing branches of the family, and it is expected that half of these will show the disease, and that the other half will be healthy. The expected proportion is, therefore, in round numbers 182 : 182. The actual result is 135 : 229, 88 THE MENDEL JOURNAL or thirty-seven per cent. of night-blind persons instead of fifty per cent. Too much stress must not be laid upon this numerical difference. It must be remembered that segregation and gametic purity can exist notwithstanding that the proportions do not accurately accord with expectation. Factors of which we have no knowledge may be at work, dis- turbing these proportions in some families or genera- tions, and leaving them undisturbed in others. No one with any real experience of living organisms, including human nature, doubts it is a fact which should be constantly held in view that living gametes within a living zygote are not marbles in a bag. The latter can be shaken up to ensure an uniform dis- tribution of the black and white ones, but gametes and human nature cannot be so treated ; we must accept them as Nature listeth they shall be. And this particular case is instructive, for there are good reasons to believe that the number of abnormals recorded does not represent the true number. The chief of these reasons arises from the promptings of a natural human weakness, 7.e., a reluctance to admit the existence of physical defects, and additionally there is upon the part of the women a very laudable desire not to jeopardise their chances of becoming wives and mothers. For the learned Curé who is familiar with the living descendants of this stock, and who has contributed to the construction of the pedigree, says: “It is considered as prejudicial to the establishment of children” to be afflicted with this disease, ‘‘ and it is therefore apt to be concealed.” VIRILE SENTIMENT 89 There is, indeed, one somewhat remarkable case where a woman kept her husband in ignorance of her condition for twenty years. Human nature being what it is, we must expect some deficiency in the numbers of the afflicted in this and other pedigrees. The next case that I have to deal with is one for which I am indebted to the kindness of Dr. Drinkwater for his permission to use. It is a case of congenital asthma. There are altogether twenty-three indi- viduals in the pedigree. The case is typically Mendelian. One individual extracted from a line in which the disease is present is free from it, and he married a normal person and had all normal children, three in number. The other extracted normal persons have not married. But four of those suffering from the disease have married normal people. The asthmatic condition is shown by the history of this pedigree to be dominant. We shall, therefore, expect in the offspring of these marriages an equal number of asthmatic and normal persons. There are, in fact, ten of each. Thus in the complete segregation of the normal and asthmatic characters, in the breeding true of the extracted recessive (normal) character, and in the proportion of the two kinds of ofispring from pairs of parents one of which is asth- matic and the other normal, the case is a definitely Mendelian one. It is of some interest to note, in passing, that of the members who have married from this family four of them (two brothers and two sisters) are affected, and one only (a brother) is normal. This is a fact of some importance to students 90 THE MENDEL JOURNAL of Eugenics, especially when it is additionally re- membered that, in regard to another disease, phthisical persons tend to marry early. It is significant because physical defects seem to be no hindrance to marriage, but apparently, in “this case, rather a recommenda- tion. The last case that I have for your consideration is one, it seems to me, of great importance, because it is very significant in many features. It not only affords us an example of hereditary transmission, but it gives us a concrete case where social sentiment and philanthropy having interfered, we can arrive at a definite judgement as to the pernicious effects of this interference. It is not an academic case that I have laboriously hunted for in the Archives of the Clinical Department of a great hospital, or in the volumes of a library. It is a case of the present time, that was reported in the Morning Post of May 25th, 1907; it is one that illustrates the prolific growth of unfitness when fed and watered by the mistaken kindness of the fit. It appeared in an appendix to the Ninth Annual Report of the Children’s Committee of the Metropolitan Asylums Board. It is written by the Medical Officer to that Board. I ask you to note that fact, because medical men are largely responsible for the birth and promulgation of the modern sentiment which we are now con- sidering. The general tone of the Report, especially the last paragraph, implies the possibility of con- verting inherent unfitness into fitness by a suitable environment. The parents in this particular case VIRILE SENTIMENT 91 demanded the return of one of their children from the Wandsworth Home of the Board, and the Medical Officer has appended the recommendation that “This is one of those cases which suggest most for- cibly the advisability of having the control of children of this mental condition for a fairly long period.” And he adds: “It will be interesting to follow his case 1n the future, and see whether his mental con- dition improves or deteriorates.”” Why does the Medical Officer recommend that this child shall be under his control for a “ fairly long period” ? Is it expected that any control, however prolonged, is goiig to transform the processes of heredity, or turn aside by a hair’s-breadth the immutable processes of Nature? Let us examine the case and see how futile is the hope. The father of this family had phthisis, and was insane; the mother was laid up for four months before the boy’s birth with spasmodic paralysis, and alterwards lost the use of her legs for some months. The paternal grandfather died of phthisis, and the paternal grandmother, who 1s still alive, is in an asylum with mania and religious fancies. The maternal grandmother died of consumption (phthisis). Nothing appears to be known of the maternal grandiather, but, since the mother suffered from paralysis, it may be that it was transmitted from her father. Now let us examine the children of these parents, first pretacing our statement with the remark that menin- gitis in this case is a form of phthisis not attacking the lungs, but the membranes of the brain. No. 1, 92 THE MENDEL JOURNAL a son, is said to be strong. No. 2, a daughter, suffers from meningitis, epilepsy, and tubercular (phthisical) knee joints. No. 3, a son, is afflicted with severe headaches. No. 4, a son, suffered a severe nervous illness, the nature of which is unknown. No. 5, a son, has had meningitis and paralysis, and is mentally dull. No. 6, a son, has had St. Vitus’ dance. No. 7, a daughter, has sufferea from meningitis, and is very irritable and subject to headaches. No. 8 (whether son or daughter is not stated) has suffered from paralysis and other nervous diseases. Nothing appears to be recorded of No. 9. No. 10, a son, is dead, and had paralysis in the legs. No. 6, the boy who is afflicted with St. Vitus’ dance, is the one whom it is recommended should be cared for out of public monies, by public officials, mn public institutions. Now we are chiefly interested for the moment in the Mendelian aspect of the heritage of this family. If we take a general view of the case we shall notice that we can reduce the pathological factors in the family history to two, if we express them in a general way. There is the tubercular taint manifested by phthisis, tubercular knee-joint, and meningitis. There is the nervous lesion indicated by mania, religious fancies, St. Vitus’ dance, severe headaches, paralysis, and epilepsy. We may therefore say that we are dealing from the Mendelian standpoint with two pairs of alternative characters, 7.e., the tubercular condition and physical normality as one pair, and the nervous lesion and nervous normality as the other. VIRILE SENTIMENT 93 A possible Mendelian prediction for such a case is that four kinds of individuals will be present in the offspring. We may state them in the following manner : 1. Suffermg from tubercular taint and nervous lesion. 2. Suffering from tubercular taint, but no nervous lesion. 3. Suffering from nervous lesion, but no tuber- cular taint. 4. Normal in both features. And the proportions in which these four types may be expected are, in the order in which they are given : 94+343+1. That is, in a family of sixteen indi- viduals, only one normal person is expected! And his normality will not be better than the normality of his race. In this family of ten only one is normal ; the others all indicate some nervous lesion, or mani- fest both a tubercular condition and a nervous lesion. And thus far this case shows a resemblance to a Mendelian mode of inheritance. There is segregation manifested in the case of the normal son, in the pro- duction of individuals showing only nervous symp- toms without tubercular ones, and in the appearance of both tubercular and nervous pathological features together in other individuals. But in a complex case of this kind many factors must be considered, and a larger number of families examined, before any degree of positive statement is justifiable. It is sufficient for the present that there is some indication of a Mendelian mode of inheritance, and that in other 94 THE MENDEL JOURNAL human characters this mode has been clearly shown to exist. Before we leave this case let us consider its social aspect. The fact of the hereditary trans- mission of the defects of this family is perfectly clear. That, at any rate, admits of no dispute. There is not an environment on earth that can make this stock other than what it 1s. Every effort that is made to save it, or to prolong it, so that the period of procreation of its individuals may be reached and lengthened out, is an effort which results in an increase of persons to whom life is a miserable and unclean burden. Every penny which is wrested from the earnings and productions of fitter citizens in order to rear an army of officials and palatial edifices for the maintenance of these unfit is by that amount reducing the number of persons to whom life means more or less of happiness and contentment. It cannot be otherwise. The penny cannot be spent upon both the unfit and the fit, any more than we can “ eat our pudding and have it too.” And it is interesting to note that what the child thinks of the pudding the philanthropist believes of the penny. Both suffer from a delusion, but the child is soon undeceived. His is a concrete matter, demonstrated in a few minutes. The philanthropist’s problem is more superficially complex, and the avenging hand does not strike until as a warning it comes too late, because with its warning it also brings destruction. Let us remember that in this family the paternal grandmother is still alive, and that the parents have VIRILE SENTIMENT 95 ten children. These two facts are important, because they demonstrate that physical and mental unfitness and civic uselessness are not necessarily accompanied by infertility, nor low fertility, nor shortlivedness. Indeed, there is here high fertility and longevity. When we bear in mind that there is good reason to believe that fertility is a character which is heredi- tarily transmitted, we have to recall the possibility that the grandparents on both sides may have ten children, that is, twenty altogether. And if these twenty marry, as the unfit always do, and they marry their “likes,” as they also nearly always do, and if each of these twenty pairs of parents have ten chil- dren, we are faced with the appalling possibility that from this family alone there are living in our present generation two hundred individuals carrying or mani- festing the tubercular, paralytic, and epileptic diatheses! And if we go forward to the next generation there may be two thousand of them, and in the third generation twenty thousand of them! An army of epileptics, paralytics, and tuberculates ! It may be asserted that I am indulging in wild and sensational alarms, and that this sort of thing has never occurred and can never happen. It is so easy for any nation to play the part of the traditional ostrich, for the sands of self-delusion are loose and soft. And it is hard to be brave when problems that can be put off for some one else to solve call upon us to solve them now, before they become so bad that their remedy must be heroic. They who assert that these things can never happen, and that a great army 96 THE MENDEL JOURNAL of useless citizens cannot be reared in a short time, have either never troubled to ascertain the facts or have ignored them. There is the well-known German case, for instance. In this example seven hundred and nine descendants of a particular woman could be traced. She was a drunkard, a tramp, and a thief. Of her descendants, seventy-six were convicts, seven were murderers, one hundred and eighty-one of her female descendants were prostitutes, one hundred and forty-two were tramps and beggars, while sixty- four lived on charity and one hundred and six were illegitimate. In the course of three generations this woman and her descendants cost the State, in trials, prisons, and workhouses, a quarter of a million of money! There is another case of a like nature, from Canada. On the Upper Hudson some generations ago there existed a child of the “ gutter type.” She ultimately became a “ prolific mother of a prolific race.” Besides a large army of idiots, imbeciles, drunkards, paupers, and prostitutes, the country records show two hundred of her descendants who have been criminals. - And this is all that our philanthropy and modern social sentiment achieves! Itrears great armies of unfit individuals, and it discourages through taxation and in other ways the increase of the fit. While the use- less are pampered the useful are harassed. While it is becoming increasingly easy and profitable to be unfit and yet live, it is becoming proportionately difficult to be fit and yet enjoy the benefits of our fitness. Freedom and licence are being granted to VIRILE SENTIMENT oF the lower classes, and liberty is being restricted for the individuals of the classes above. The whole-influence of this modern sentiment is trending in the wrong direction. It sets out with the belief that it can effect the salvation of the unfit by improving their environment. It will end by achiev- ing the destruction of the fit, without having even gained the salvation of the unfit. It commenced its operations by creating an army of grand-maternal officials to guide the footsteps of the weak, the stupid, the inane, the imbeciles, and the physical and moral wrecks. It will end—indeed, it has begun to end—- by the creation of an army of lower-grade officials who are rapidly degenerating into petty tyrants, who seek to restrict the successes of better citizens that have achieved what they could not, and have excited alike their cupidity and their vanity. And this, it appears to me, is the social aspect which the case of the Metropolitan Asylums Board presents to us. Through- out history every effort to save the unworthy has reacted to the detriment of the worthy. It is time we looked more seriously both at the teachings of History and Biology. A popular impression exists that by marriage it it possible to minimise in the next generation the evil qualities of one of the two partners by the good qualities of the other. It is a belief which is based impliedly upon the conception that underlies what is termed blended inheritance. Its faith is rooted in the assumption that the offspring represent a blend of their two parents, and of their more distant ancestry, G 98 THE MENDEL JOURNAL too, through the channel of their parents. It is implied that the evil propensities of the one will be diluted, in the processes of heredity, by the goodness of the other. It seems, however, to be forgotten that, even if it be true, the goodness is diluted too. But if the processes of segregation and gametic purity are operating in Man, there is an end of this belief. For no marriage of fit with unfit, of virtue with vice, of truth with falsehood, of courage with cowardice, and of physical vigour with physical weakness will result in any diminution of the alterna- tive bad qualities. Marriage will merely procreate and multiply the individuals who manifest them. When Friedrich Nietzsche, in his “Thus Spake Zarathustra,’ wrote the three following paragraphs in the chapter “Of Child and Marriage,” he was nearer than he imagined to the truth, and he antici- pated, in a few words, the EKugenic doctrines of the present :— “Worthy and ripe for the significance of earth appeared this man unto me, but when I saw his wite earth seemed unto me a madhouse.” “Yea, I wish the earth would tremble in convul- sions whenever a saint and a goose couple.” “This one went out for truths like a hero, and at last secured a little dressed-up lie. He calleth it his marriage.” The follies and impulses of youth are proverbial ; even the youth who ultimately may blossominto genius or to a high degree of civic worth, and in his maturity may aspire to high ideals, is not free from the errors VIRILE SENTIMENT 99 incidental to his age and temperament. Students of the Divorce Courts are familiar with examples that serve to illustrate the disastrous consequences that as often as not follow in the train of early folly. Whena youthful member of a noble family, forgetting, in his ardent impulses, the traditions and glories of his stock, marries a ballet girl; or when a young man of impulsive moods but of good parts otherwise, and descended from a family of good social position, and who will later develope all the aspirations that belong to his social class, accepts a partner without any solid qualifications and who has temporarily ascended from a lower social level, and will manifest the instincts and inherent defects of her class; or when a young man, who because of the stock from which he springs, will later on in life become a more serious citizen, marries .a “ dressed-up doll”; or when a woman, yet in the vernal stage of life and of refined instincts, consents under parental pressure to marry a rich man of coarse habits whose years are strewn with the sere and yellow leaves, no Heaven-made conception of the nature of human marriages can avert the inevitable consequences. It is a serious question for the student of Kugenics whether a youthful error of judgement in marriage, by which “ a dressed-up lie ” and “ a goose ’ is chosen as a mate, should be visited with permanent and lifelong punishment. The time has passed when the dogma of “ heaven-made marriages” should be allowed to operate to the detriment of much that is best or good in the community. No marriage that declares by its results its obvious inequalities and > 100 THE MENDEL JOURNAL disastrous disabilities should be permanently main- tained. If one of the two partners is capable and the other grossly incapable, if one is intelligent and thrifty and guided by high ideals, and the other is foolish, thriftless, and incapable of rising above the lowest aspirations, then the marriage, as in a part of the days of Rome, should be possible of mutual dissolu- tion. For to permanently bind a good citizen to a bad one is waste of the good civic material. I am aware that the problem is a complex one, and it is not simplified by the possibility that the incapacity to make a good judgement in the choice of a mate may be an inherent quality, and not always a passing aberration of youth, and will be therefore hereditarily transmitted. But in spite of that, perhaps it would be wiser to give an individual at least one opportunity of rectifying a youthful mis- , judgement. I am treading, I know, upon dangerous ground, and I hope that I shall not be misunderstood. I trust that nothing which I have said can be construed into implying any sympathy with those ideas of pro- miscuous marriage that in certain quarters are being preached by men whose fanaticism or desire for notoriety is greater than their knowledge of either the history or nature of mankind. I, for one, repudiate all such ideas. But the sanctity and holiness of marriage may be destroyed when earthly considera- tions and ideals are ignored by a dogma that has no justification in true belief or in truth.* * See Appendix, pp. 113 and 114, VIRILE SENTIMENT 101 If now we take a general survey of all the cases which we have considered, we shall agree, I think, that the evidence is pretty clear in regard to some of the qualities of man that there is segregation in their hereditary transmission from generation to genera- tion. There is reason to believe that future investi- gation will widen the number of human characters which so behave. There is no reason to believe that the mental and moral qualities of man are trans- mitted in any way different from the physical characters. Indeed, the evidence indicates that they are hereditarily handed down in the same way that the physical ones are. Without citing more recently- investigated cases, it will suffice to pomt out that it is a historically-known fact that the eloquence of Marcus Antonious was a hereditary gift, and that the high ability of Brutus’ family extended through many generations, and was manifested in the dis- charge of the duties of the many great posts which that family occupied during successive genera- tions. If, then, it shall be ultimately shown that segre- gation of alternative characters and gametic purity is the rule of man’s hereditary processes—if the clear indications of the limited cases already known become a generalised expression of a general process—and justify us in saying that segregation and gametic purity express a natural law of man’s heredity, what bearing will this have upon all social questions and upon questions of even wider moment ? Looking over the cases which we have already 102 THE MENDEL JOURNAL considered, and substituting for such physical characters as short fingers, curly hair, and stationary night-blindness certain bad civic qualities such as those which characterise the loafer, the wastrel, the congenital drunkard, the habitual criminal, the con- genitally tuberculous, the mentally deficient, the thriitless and generally incapable persons, and others, how will the problem work out? When we recall with what persistency the peculiarity of stationary night-blindness and_ short-fingerness passed on through the generations—passed on for two hundred and seventy years in the case of the former character, and in the latter case for about one hundred and eighty-nine years—without any known alteration, it is time that we began to consider what types of citizens our philanthropy and charity are breeding. It is time we faced the problem. It is time we learned how little environment can do, and how much the inborn qualities determine all for us. In the multitudinous efforts which are being made for social reformation it is to be hoped that the beauties of character will not be forgotten, and that the road to success will not be made too easy, nor the road to destruction too difficult. If by our social efforts we are breeding, rearing, and accumulating an un- desirable stock that cannot or will not work, that cannot or will not tend its children, that cannot or will not be sober and thrifty, that cannot or will not educate and feed itself, that cannot or will not attaim an intellectual and moral level that is necessary for social life, then we are advancing along the road that VIRILE SENTIMENT 108 brings destruction to the fit in order that the salvation of the unfit may be attempted. There can, I think, be no doubt that those circum- stances which are attendant upon the workings of a democratic state of society tend to create and to fan into larger size a flame of emotional sentiment that becomes wildly excited over the imagined or exag- gerated suffermgs of the weak, incapable, and generally unfit. We have only to listen to the hust- ings speeches of the politicians, especially to those of the type delivered by Mr. Winston Churchill and Mr. Lloyd-George, to see how pernicious are the workings of democracy when the leaders are unworthy of their position. The same type of morbid sentiment and craven fear that gave the mob of Rome its free circuses and free bread, that gave to the Athenian multitude expensive theatrical shows free of cost to themselves, and placed needy adventurers and profligate dema- gogues of the type of Chares in positions of executive and legislative power, is to-day in our own community increasing the privileges and rights, and decreasing the civic responsibilities of the lower classes of society, and simultaneously is placing power in the hands of men who either represent these classes or are sprung from them. Greece and Rome possibly fell in ignor- ance. There may have been none among their counsellors who saw the importance of the facts of heredity, or, even more probably, did not recognise that such facts were part of the operations of Nature. If England has reached her zenith it cannot be said 104 THE MENDEL JOURNAL that her decline is due to ignorance. Rather, it must be said that it is comcident with a period of great activity and advance in biological knowledge. To- day the truth of Evolution, the Processes of the Survival of the Fittest, and the fundamental facts of Heredity are clearly grasped and understood by biologists. At no period in human history has knowledge been so extensive, so accurate and so carefully generalised as now. No one acquainted with human life and with lower animal life doubts the identity of the natural processes that act upon both. Man’s ethical and esthetic qualities are not new creations—only elaborations of qualities present in the living kingdom long before he came. And beneath his ethical and esthetic nature lies the animal. In the long run the animal instincts, hunger and pro- creation, determine man’s conduct. We need only recall the French Revolutions to remind us of the fact. Let us not forget the guillotining until the headsmen sank worn out ; followed by the fusillading of little children—* the wolflings”’ of Marat, “ who might grow to be wolves ”—and of women with chil- dren at the breast ; of wholesale drownings, of women stripped naked before they were drowned, and of mothers dragged to the guillotine to witness the slaughter of their innocent children. Truly did Carlyle say, ‘“‘ Cruel is the panther of the woods, the she-bear bereaved of her whelps ; but there is in man a hatred crueller than that.” The angelic gloss on man is but a surface-coat painted in times of peace and prosperity, but cast off in times of distress and VIRILE SENTIMENT 105 hunger. We must not forget the recorded facts of ship-wrecked and starving men casting lots for their next meal, and feeding upon him whom chance had marked. Neither must we forget the County Courts, nor the now historical episodes of the Marshalsea. “Give me the pound of flesh which is due to me, or die for aught I shall care,” is the general formula that designates the doings of mankind, even in normal times. Ido not regret that itis so. Ido not, in even small measure, denounce Shylock. I have come to recognise that the weak demand the “pound of flesh” from the strong whenever they can, no less than the strong demand it of the weak. Be careful, therefore, philanthropists and social sentimentalists, that in your frantic haste to procreate and preserve the civically unfit you are not bringing into being a 74 great herd whose demand for its “‘ pound of flesh ” shall be none the less emphatic because it cannot earn the value of that pound, or, having earned it, finds that others, stronger and fitter, have already consumed it. It is a dangerous problem, this of the multiplication of organisms, with that philogamic passion at their rear ever unresisted, driving onwards. It is dangerous even if the multiplication is that of the fittest. But if for a while you shall aid the maintenance and increase of the unfit and unfittest it means ultimately an annihilation on a scale passing beyond all comprehension and transcending all con- ception. It is therefore wise to heed how far the hour may be distant when by the voice of the great shipwrecked herd the casting of the lot may be 106 THE MENDEL JOURNAL demanded, and in spite of your philanthropy and mistaken kindness you shall serve as food, to stay for a moment the hunger of that wretched and helpless mob that your misguided sentiments called into a larger existence. For, as Vergniaud said of the execution of the Girondins: ‘‘ The Revolution, like Saturn, is devouring its own children,” so it may be said of your sentiment that it will devour you and yours. If, in defiance of the teachings of biological science, modern sentiment persists in breeding and rearing a helpless race that cannot by its own efforts satisfy its hunger, or if it will persist in rearing a race that mani- fests criminal and brutal instincts; if, in order that the lower democracy may be appeased, the unfit, the sluggish, the lazy-rapacious, and the cunning demagogues are in turn extolled, bribed, and main- tained in life at the expense of the fit and worthy, then England, in spite of her greater knowledge, has commenced to follow the path that led to the destruc- tion of Rome and Greece. If the ruling classes of this country, the Aristoc- racy, have forgotten their duties, which is to face and enforce the truth of their generation, or have lost their courage, then it must fall to the more vigorous and less esthetic portion of the Intellectual Classes to take their place. The danger is clear and eminent enough. With the Lower Classes increasing their birth-rate, while the Middle and Upper Classes are remaining stationary ; with certain well-intentioned but mistaken members of the higher social classes VIRILE SENTIMENT 107 making remarkable efforts at much personal sacrifice to lower the infantile mortality of the lowest strata of society ; and with workhouses, infirmaries, and lying-in hospitals using baby incubators to fan back into existence the weakly lives that benign Nature, in the noblest interests of her race, demands shall perish, there is no need of a vivid or expansive imagination to picture the end of a nation that has become thus far decadent. It seems that we, as a nation which have so long professed a belief in a merciful and loving God, have forgotten the old cry of resignation, “* Thy will be done,” that brought comfort, courage, and virility to Englishmen of the past when the processes of Nature eliminated their weak and sickly. They understood not these processes, and in ignorance resigned themselves in a noble faith that all was work- ing towards a beneficent end; and, in their resigna- tion, found their salvation and that of their race. . The generation of to-day believes it understands these processes, and in the arrogance of its belief has played the coward and traitor to its heritage. In its senti- mental interference with the workings of Nature it is leading the community to its destruction. In its attempts to be superhuman it has become inhumane. In its efforts to save life it has increased the number of individuals to whom existence means misery and pain, and is decreasing those to whom life means health, happiness, and progress. It recalls to my mind some of Wordsworth’s lines in “ The Excursion ” :— “ Vain-glorious Generation! what new powers On you have been conferred? What gifts, withheld From your progenitors, have ye received, 108 THE MENDEL JOURNAL Fit recompense of a new desert? What claim Are ye prepared to urge, that my decrees For you should undergo a sudden change ; And the weak functions of one busy day, Reclaiming and extirpating, perform What all the slowly-moving years of time, With their united force, have left undone ? By Nature’s gradual processes be taught ; By story be confounded! Ye aspire Rashly, to fall once more; and that false fruit, Which, to your over-weening spirits, yields Hope of a flight celestial, will produce Misery and shame. But wisdom of her sons Shall not the less, though late, be justified.” The generation in which we are living seems to be fond of poetry of an emotional kind. Let me address to it a few more lines of a type of poetry which is less sentimental, but more truthful :— “The Moving Finger writes: and, having writ, Moves on: nor all your Piety nor Wit Shall lure it back to cancel half a Line, Nor all your Tears wash out a Word of it.” It would be well if we endeavoured to understand the real significance of these lines in relation to the doctrines of modern sentiment. In more than one way it points to us the irrevocable consequences of- every attempt that is made to interfere with the beneficent workings of Nature. The “ Moving Finger writes,” and then ‘“‘ Moves on,” is but a poetical way of warning us that for every committed act, social as well as individual, there are both immediate and multiplied remoter consequences,-the misery entailed by which cannot save us from their inflexible opera- tion, nor all the tears engendered by them wash out a single punishment which they inflict. We all remember as children reading of the traditional King Canute, who, at the instigation of his flattering courtiers, sat on the sea-beach and bade VIRILE SENTIMENT 109 the tide recede. He bade the immutable processes of Nature that determine how and when and to what extent the tides shall rise and fall to cease. But no decree of man or king can alter by the fraction of an inch the rising of the tides. It would be well if we recognised that human life is controlled and deter- mined by processes that are as immutable and as merciless as those which govern the tides or determine the movements of the planets. We can learn to understand them and to obey them, but we cannot destroy or diminish them. Unfortunately there exists to-day a Canute who is not traditional, but figurative. We may regard him as the symbolic emblem of modern sentiment. He sits upon the sea-shore of current deeds, and, looking forth upon the rising events of Futurity, demands that they shall cease to rise! The hour of his disillusionment is not far off ; it is nearer than he imagines, for the shore upon which he sits shelves less than he believes. By not the fraction of a second will the. tide of Nature’s processes be delayed or hastened in its rise upon the Future of the Nation. What we sowed yesterday is to-day germinating, and will be reaped to-morrow. There is time, even now, to dig up the sprouting tares before their harvest shall have destroyed the wheat. Who, with hoe and rake in hand, will go forth and lead the way ? Will the natural rulers of the Nation, the Aristocracy, do it, or must the more vigorous and virile of the Intellectual Classes—the Upper Middle Strata of Society—ignoring its poetically esthetical section, usurp their position and discharge their functions ? 110 THE MENDEL JOURNAL APEENDEX. Added February, 1909. Note to p. 68.—By “ Genetics ” is understood that branch of Biology which studies the phenomena of heredity. Quite recently, owing in part to a redis- covery of Mendel’s generalisation and in part to the large and increasing volume of experimental evidence in corroboration of it, the study of heredity has now for ever emerged from the chaos which marked its previous condition. An appeal to accurately con- ducted experiments is the soundest and safest method by which the problems of heredity can be solved. It may be urged, indeed has often been urged, that we cannot apply the experimental method to man. But so far as he is concerned, experiments on a vast scale have already been performed. It is not the want of experiment that we lack, but the proper and accurate recording of the results. Blue eye with blue eye, brown eye with brown eye, and brown eye with blue eye have been mated together in numbers that dwarf our most colossal experiments—even those of Prof. Bateson with an offspring of twelve thousand five hundred chicks—and yet the only accurate analysis and record of these facts which has been made is that by Mr. Hurst of quite recent date. North American Indians and negroes have been crossed with Europeans many hundreds of times and their progeny have intermarried and _ been married back to the original European or coloured stock still more frequently, and yet nowhere is there VIRILE SENTIMENT 111 to be found a scientifically recorded statement of the facts. The material exists, but 1t wants scientific analysis and recording. If this address should be read by any persons of leisure, or by medical men, who may at any time be resident in Canada, and who may become acquainted with families where intermarriage between Europeans and Red Indians has occurred, they may render services of the greatest value by taking photographs, both full-face and side-view, of the grandparents, parents, offspring, and of as many collaterals as possible, and recording as carefully as can be the colour of the skin (both that exposed and that protected), the colour of the eyes, and the colour and nature of the hair of each individual. More valuable still would it be if a lock of the hair of as many persons as possible were obtained. If it is not possible to obtain complete pedigrees, then partial ones will serve the purpose, provided that the nature (whether half-castes or otherwise) of the different individuals is accurately ascertained. A letter addressed to me at the London Hospital Medical College, London, E., will always reach me. In the same way marriages between negroes and Kuropeans and the various marriages of the mulattoes could be recorded. In the case of mulattoes, quadroons, and octoroons it is desirable, if possible, that observations as to differences in tints in each class should be recorded. It is also very essential that not only skin colour, but hair colour, hair texture (whether European or woolly), and eye colour should be recorded. Other facial features will, of course, be recorded in the photo- 112 THE MENDEL JOURNAL graphs. The results of crossing albino negroes with normally pigmented members is also urgently required. The same considerations hold with regard to human albinism generally. Much of the published records are incomplete and not precise enough. The case of Josephine Chassot described on p. 77 is an example of this. She is described in the original 4 publication as “affected with albinism,” as having “pink eyes (pupils),” but “with peculiar lilac- 6 coloured irides,” and as having a “ very white skin.” But nothing is said about her hair colour. When described in comparison with her sister, who is stated to be dark, she is said to be fair (blonde). Such a description may mean that her hair is flaxen or white. The other albinoes in this pedigree are simply des- cribed as “albinoes,” or as being “affected with albinism.” But as to whether they are complete albinoes having no pigment at all, or are so-called partial albinoes having flaxen hair, we are left to infer. Note to p. 68.—A definition of “ Kugenics ” by its author, Mr. Francis Galton, may be found in “ Nature,” Vol. LXX., p. 82,1904. “ Hugenics is the science which deals with all influences that improve and develope the inborn qualities of a race.” The essential basis upon which “ Eugenics” is founded exists in the consideration of certain postulates which all will accept, z.e., “that it is better to be healthy than sick, vigorous than weak, well-fitted than ill- fitted for their part in life. In short, that it is better to be good rather than bad specimens of their kind, VIRILE SENTIMENT 113 whatever that kind might be.” “The aim of Eugenics is to represent each class or sect by its best specimens, causing them to contribute more than their propor- tion to the next generation ; that done, to leave them to work out their common civilisation in their own way.” Mr. Galton thinks that learned and active societies desirous of promoting Eugenics might proceed by the following methods: “ (1) Dissemination of a know- ledge of the laws of heredity so far as they are surely known, and promotion of their further study. (2) Historical enquiry into the rates with which the various classes of society (classified according to civic usefulness) have contributed to the population at various times, in ancient and modern nations.” Mr. Galton then adds: ‘‘ There is strong reason for believ- ing that national rise and decline are closely con- nected with this influence.” ‘‘ (3) Systematic collec- tion of facts showing the circumstances in which . large and thriving families have most frequently originated. (4) Influences affecting marriage. (5) Persistence in setting forth the national importance of Kugenics.” With reference to the third and fourth of these proposals—and which may also partake of the nature of a note to pp. 99 and 100—I would like to add that I have for some time now been endeavouring to collect facts bearing on the causes of what we may call “unhappy or incompatible marriages,’ and I am desirous of obtaining further information from those who are in a position to give it. Names are not H 114 THE MENDEL JOURNAL necessary, and all information will be regarded as confidential. There is reason to believe that in some cases these marriages are the result of an inherent incapacity upon the part of one of the two partners to make a felicitous choice. | An inherent incapacity of this sort would, therefore, because it is inherent or congenital, run through members of the same family, and would be manifested by that family having a larger proportion than usual of such unhappy marriages. If any one of my readers is acquainted with families of this kind, I shall be glad if they will communicate with me. In other cases, my present information suggests that these marriages are the result of a too early union. Ata certain age, say, from twenty to twenty- five, a pair of individuals may possess temperaments and capacities that are more or less compatible and could harmoniously exist together. But at a certain later age, let us say about twenty-six to thirty, one or both individuals undergo a change in temperament, character, and ideals, perhaps as divergently as it is possible to conceive, and the marital state becomes incompatible and impossible. In this connection, it is a fact of some interest, which I have no doubt others have observed, though perhaps they may have given it no more than passing consideration, that there exist people who instead of becoming wiser with experience, become more foolish, and who instead of becoming more proficient become less so, as they pass on towards their primal years. If such per- sons passably discreet, tactful, and proficient at the VIRILE SENTIMENT 115 time of marriage, for the positions which their youth- ful years are likely to occupy, are married to partners who improve in all qualities as time advances, it is clear that the barrier of incompatibility must widen with the passage of each year. I do not suppose that these cases are very numerous, but still they exist. And it seems to me a very important matter, because I think that these changes of which I speak are inherent or inborn in their nature, and can no more be avoided than the appearance of the antlers and of the fighting instincts of the stag when it has reached its maturity ; and cannot be commanded to stay their appearance any more than can _ those sexual instincts and secondary sexual characters which inevitably appear in human beings at a certain stage in their developement. This appearance of new characters and new qualities at different stages in the lives of individuals, and of one kind for one person and of another kind for another, is of far wider social import than its application to married life. Its very existence vitiates a great deal of the medical evidence that was given before the Committee on Physical Deterioration and Degeneration. This latter question is not so much a medical one as it is a biological one, and the medical evidence, though valuable, is not the most important that can be adduced, and, in fact, in many of its conclusions it is erroneous and in its nature entirely misleading. Note to pp. 102, 104, and 106.—While this address has been passing through the press, four events of some 116 THE MENDEL JOURNAL significance have happened. One of these is the earthquake of Messina and its attendant incidents. Among these incidents there is one which is, in history, of such ordinary occurrence that in our familiarity with it we treat it with an altogether un- deserved indifference. I allude to the fact that among the stricken and starving inhabitants, “ fierce fights with knives for bread” occurred. And, after some seeming order had been restored, the distribu- tion of food by the authorities took place in the pres- ence and under the protection of an armed guard. Where hunger presses and it cannot be satisfied, blood will inevitably flow. The second incident is the occurrence of a fire at a cinematograph exhibition in the East End of London. It was apparently quite a simple and not very dangerous affair. The cinematograph film caught fire and .made a flare altogether out of proportion to its possibilities of danger. According to the newspaper reports of the operator’s statement of the accident, the cinemato- graph machine was enclosed in an iron box. None the less, a panic seized the audience, and the women, in their frantic endeavours to save themselves, trod down the children, and the men trod down both women and children in their equally frantic efforts to save their threatened lives. . “ Each one for him- self and destruction to the hindmost,’” expresses this event. And within the past two years there have been a succession of similar events. The third incident is a “ nine-days’ wonder.” It VIRILE SENTIMENT 117 ought rather to be an iconoclastic fact ever fresh in our memories. Whenever it is recalled it will probably be spoken of as the “ Tottenham slaughter.” Posterity quickly forgets. In a twelve-months’ time the affair will have sunk into oblivion, in company with a great many more unpleasant facts that ought likewise to be remembered. Let me, therefore, record its essential details. A Russian outlaw had sought the hospitality and protection of English laws. He had lived in this country apparently for some months, but for a fortnight of this time he had been employed in an English factory at Tottenham, but gave up the work “ because it was too hard.” During the interval he had learned that the wages for the employés were brought from the bank in a motor-car at half-past eleven o’clock every Saturday morning. On the Saturday morning of January, the 23rd, 1909, this outlaw, in company with a companion, who was similarly an Anarchist outlaw from Russia, waited at the appropriate hour outside the entrance to the works. Both men were armed with revolvers and with pocketfuls of cartridges. Their intention, there- fore, was quite clear and premeditated. As soon as the messenger had alighted from the car with the bag of money, it was seized by one of the desperadoes, and in the struggle which ensued the messenger was fired at and wounded. ‘The assassins then made off, and endeavoured to clear a path of escape for them- selves by indiscriminate and wild firing, right and left of them. The hue and cry was raised, and chase was given. Some of the pursuers were on foot, some 118 THE MENDEL JOURNAL on horseback, and some in a motor-car. The pur- sued were chased for a distance of five miles, making their way towards Epping Forest. In the course of their reckless flight, they boarded an electric tramway, fired at and wounded some of the passengers, and one of them compelled the driver to proceed at a rapid pace by holding a revolver at his head. While this one was thus engaged at the front of the car, the other was at the rear, and in an indiscriminate fashion was firing right and left. At a certain point they left the car, which they then believed to be nearing a police- station, and boarded a milk-cart, which they drove in another direction at a furious pace. This they ultimately left, and then took to the fields. Soon alter this one of the two was brought to bay, and he shot himself, but not mortally; he was taken prisoner, and subsequently died from meningitis in a hospital. The other effected his escape by an almost superhuman effort, and finally took refuge in a small house. Here he was at last shot by one of the police- men who had taken part in the chase and who had borrowed a revolver. In the course of this bloody fracas, a policeman and a small boy were shot dead and twenty-three persons were wounded. One important moral we may draw from this event: The civically unfit under an autocratic régime are similarly unfit under a democratic one, even though it be pervaded with a benevolent sentiment. But the autocratic régime knows how to get rid of them, and the democratic one how to receive them. VIRILE SENTIMENT 119 Here, then, in our midst, modern sentiment allows a type of criminal not only to exist, but also to breed ; and this type is one so devoid of all sense of civic responsibility and so destitute of the rudiments of social instinct, that the individuals which constitute it are deliberately prepared to ruthlessly and indis- criminately murder all who shall endeavour to frus- trate their criminal desires; and even children are not excluded. Do not let us delude ourselves by believing that only Russian Anarchists can perpetrate these deeds. This type of criminal is a mutation common to all nations and to all races of mankind. They are not the product of any social conditions any more than the short-legged and long-bodied ram which gave rise to the Ancon race of sheep was the product of special farm conditions. They are germinal mutations, and as such will breed true to their mutant character. The fourth event occurred on Monday, the 11th of January, 1909, at half-past seven o’clock in the morning. The Democracy of France has been per- meated, in common with other Western European nations, with the spirit of sentimental regard for all types of civic unfitness which we have been con- sidering. “ And France has been paying the penalty. For, encouraged by the knowledge that this morbid sentiment would protect murderers from the guillo- - tine, and would only send them to expiate their crimes beneath the genial climes and the cocoanut trees of French Guiana, with a profusion of tobacco to smoke and of abundant leisure, a gang of desperadoes 120 THE MENDEL JOURNAL had terrorised the North of France by a great number of motiveless and ruthless murders. The leader of this band had committed two hundred and fifty such murders, and he was tranquilly playing cards and contemplating the happy hours he would spend under the cocoa-palms, when he and his companions, three in number, learnt the fact that even the stupidest sentiment must some time come to an end. For at last, craven fear had attained what common-sense could not, and these murderers were condemned to death. An enormous crowd assembled to witness the execution. Throughout the proceedings the crowd did not cease to raise shouts of “ Death, death!” The “delighted,” “‘ exultant,” and “ excited.” France has, through the agency of her sentiment, crowd was. bred her criminals and unfit, just as we are doing, and now she must meet the situation by a ruthless vengeance and wholesale destruction, just as we shall have to, or herself sink into “death vomited in great floods.” Lasser favre, which leaves all unfitness to reap its own destruction as fast as it arises, achieves the end more humanely and much quicker. I have appended these four cases in order that we may be reminded what will be the conduct of the human multitude when the day of adversity and stress arrives. Unless a nation is composed of reliant and self-supporting individuals this day of adversity is inevitable. For so soon as our morbid sentiment has reared a shipwrecked horde of helpless incapables of such dimension that the fit cannot or will not any longer bear the burden of its support, then the day of VIRILE SENTIMENT 121 reckoning will be with us, and “ all our piety and wit and knowledge will not cancel by half a second ” the casting of the die, nor all our tears, nor lamentations, nor unspeakable anguish of our womenkind and our children will avert by a single drop the streams of blood that will wash the streets of our great cities. It is not well to forget Nantes of the 14th of December, 1793. The Girondins came to regenerate a stricken France. They were men of talent, of courage, of constitutional principles, but the great impotent herd of the Sansculottes guillotined them by the verdict of a “ Patriot Jury ” of “ terminer les débats ” for their services to their country. Are we sure that we are not breeding a race of Sansculottes who will demand license for themselves and terror and destruction for every good citizen? The beauty, courage, nobleness, and unspeakable pathos of Jeanne-Marie Philipon as she appeared before the blood-besodden crowd of the lower French Democracy did not save her, nor will it, under like circumstances, save the best and highest of our women from similar bloody and blindly venge- ful deeds, when once our sentiment has called into being a multitude that is helpless to live by its own efforts and meets all resistance to its demands for the confiscation of the property of worthy citizens by bloody and ruthless executions. It does not matter whether the adversity which excites these deeds is due to the corruptions of a Royal Court, to the impotence of a brutal multitude, or is the product of an excitable and morbid sentiment. Adversity, cupidity, and hunger are not altered in their nature 122 THE MENDEL JOURNAL or their effects by the method of their origin. And the lower the social class we rear and pamper, the more animal and the more hemal will its instincts and its acts become. England, then, should beware of that fatal lethargy which rests on the belief that evil and destruction cannot visit a nation that was once— and may yet remain if she so wills it--the greatest of the recorded Ages. Upon the walls of her Temple the inscription of her doom is being written. Shall it be erased before it is completed, and in its place the motto of vinlity inscribed: ‘‘ England shall be Fit and Strong?” If that is the nation’s desire, the day of preaching is done, and the hour of deeds has come. For, on the dismembered Empires of the Past, the Warning Angel of Historical Experience stands, pointing the way to England’s safety. The admonitions that rise anew from the ruins of Greece, of Rome, and of Venice, bid us desert the road of a too widened Democracy, and turn to that which leads towards a wise, an understanding, and a broadened Oligarchy. Never to any country came the opportunity that lies at England’s feet to-day. With an Artizan Class unequalled for skill by any rival; with com- mercial organisers of great resources, subtlety, and boldness of methods; with a Middle Class eager for enterprise and gifted with productive capacity ; with a Literature studded with gems as precious as any the world contains, and the names of whose poets and prose-writers are honoured in every seat of culture VIRILE SENTIMENT 123 throughout the world; with a Scientific Hierarchy the greatest of its Age; with a Landed Gentry, the blood of whose sons has been shed in their country’s welfare on the torrid deserts, fertile plains, and frozen snows of every Continent, and whose deeds of valour, codes of honour, and conceptions of duty have furnished an example to every age that shall look back to them for inspiration ; with an Aristocracy that has succeeded beyond all others in the govern- ment of diverse peoples, England is blessed with that combination of qualities that should render her invincible against the world. And yet, mighty though she inherently is, sound in sinew, strong of intellect, and prolific of wealth, she hes prostrate before the fetich of a lower caste Democracy ; her splendour sullied, her energies paralysed, her culture forgotten, her knowledge neutralised, and her tradi- tions ignored, amid the blatant din of government by untruthful mural posters, by uncouth and pyrocephalic demagogues careless of veracity, and by newspaper placards that shamelessly and ostentatiously offer bribes to the lowest and least worthy of Englishsociety. Truly the hour and its running sands call for leaders who, like Pitt and Wellington, the Aristocrats, shall think of country before salary, of honour before the acclama- tion of a multitude, of principle before expediency, and of the preservation of the Fit, not the bribery of the Unfit ; who shall put Biology before Sentiment, and, wedding the former to History, shall build anew, with the aid of a more virile people, the foundations of a yet greater Empire. But if she is to accomplish 124 THE MENDEL JOURNAL this, then in the immediate self-denial of a mistaken sentiment and in the struggle with the errors of to-day, she must sow the seed that shall ripen into the glory and the welfare of to-morrow. The biological and historical hour-glass alike warn her, that the sands of her greatness have nearly run their course. Is England still great enough to stem the Tide and turn back the Flood? Hf so, her rejuvenated but broadened Oligarchy shall proclaim the answer. MENDELISM AND SEX. An Address delivered to the Mendel Society, 29th March, 1909. Bs THE question of the determination of sex is an old one. It is a problem which has been much debated along various lines of inquiry. Until recently it not only eluded all solution, but gave no promise of solution. Recent Mendelian experiments, commenced about the year 1900, had not, however, been very long in operation before the conclusions to which they led afforded clear suggestions that the problem of sex could be investigated by the same methods and with the aid of the same directing principles. Mendelism, in fact, has provided the key by which the question can be accurately and experimentally answered. Much however remains to be done, for the knowledge which we now possess is to be regarded as in the nature of a right beginning rather than as a final solution. Until quite recently it was generally believed that external conditions determined the sex of 126 THE MENDEL JOURNAL individuals. It was assumed that if we knew these conditions we should be able to decide at will the sex of the unborn young. Acting under the influence of such a_ belief, several investigators designed and carried into operation various experi- ments that were intended to show the part which different factors in the environment played in the determination of sex. The well-known experiment of Yung with tadpoles was of this class. In this experiment, certain tadpoles were fed on very nutritive and abundant food, while others were fed on less nutritive and limited material. It was found that there was a larger percentage of female frogs derived from the former and of male frogs from the latter. The conclusion thus suggested itself that females were determined by excessive anabolic or building up process of nutrition, and males by that antithetic process in which anabolism only just keeps in excess of the katabolic or breaking down processes. But there were several sources of error in such an experiment. The most serious one is indicated by a similar experiment with the caterpillars of certain butterflies. In this experiment it was ascertained that under-feeding did not result in the production of an excess of males, but in the elimination of the females. There being a heavy mortality among the females, it is a natural con- sequence that there appears to be an excess of males. But it is apparent and not real. -It is, therefore, in reality, not a case of sex-determination by environmental influences, but one of survival MENDELISM AND SEX 127 of the fittest. The males are more resistant to the harsh effects of a low nutritive diet than are the females. Moreover, the interesting case of the bee should have been sufficient to expose the fallacy of the belief that the environment can determine sex. The queen bee lives under special and uniform conditions, and she is fed on highly nutritive material. If conditions determine sex, then she should produce offspring definitely predominating in one direction with respect to sex, for it should be mainly feminine or mainly masculine. Moreover, if high nutrition determined the formation of females, then the parthenogenetic eggs of the unfertilised, but specially fed, queen bee should be mainly or wholly female. But the reverse is the case. The unfertilised eggs all produce male bees or drones, while females or workers are alone produced from the fertilised eggs. The simple fact that the act of fertilisation thus determined the sex of the individuals arising from fertilised eggs is by itself sufficient to show that sex is determined within the germ-cells, and is not dependent on environment. It is recognised now, by those who are engaged in the experimental investigation of the question of sex and of cognate problems, that sex itself, like other qualities, is predestined in the germ-cells, and is definitely determined by fertilisation. And, when it has thus been predestined and determined, external influences cannot alter it. It is, however, possible that in particular cases the proportions in which the sexes may appear are determined by 128 THE MENDEL JOURNAL environmental influences. In the case of the butterfly caterpillars mentioned above, certain conditions associated with semi-starvation, while they do not determine sex itself, do determine the proportion of males and females, by unduly eliminating the latter without influencing the former. But this difference in the resisting powers of the two sexes is probably exceptional, and, as a rule, the sexes are similar in their inherent responsive powers towards special environmental conditions. In the majority of cases, where we are in possession of sufficiently accurate statistics, it appears that as a general fact the sexes are produced in equal numbers. This is so for Man, for the lower animals, and for unisexual plants. The production of the two sexes in equal numbers is a_ significant fact from the Mendelian standpoint. It had no meaning whatever in pre-Mendelian days; but now its interpretation is clear. The individual distinct- ness of the sexes is also a significant fact. It indicates the complete segregation of maleness and femaleness. This segregation of the sexes and their occurrence in equal numbers at once suggested the well-known Mendelian ratio of 1:1. This ratio is the result of mating a Mendelian hybrid* with an individual carrying the recessive character, and it indicates that one of the two sexes is a dominant to the other. If we use the symbols which have been previously described,* and if we tentatively regard the female as a dominant hybrid and the * For definition and explanation see pages 63 and 67. MENDELISM AND SEX 129 male as a pure recessive, or vice versd, then they will be respectively symbolised as DR and RR. A reference to the table of matings on page 69 will show that the expected offspring from a D R parent mated with an R R one, will consist in equal numbers of D R’s, which in this case will be females—if we regard the female character as dominant—and of RR’s, which will be males. Thus far, then, the two general facts already known to us, namely, the segregation and numerical equality of the sexes, strongly suggest that sex is predestined in the germ- cells and is hereditarily transmitted in accordance with the Mendelian principles of gametic purity and segregation. It was not however until quite recently that experi- ments specifically designed to answer the questions presented by sex have, by their results, extended the suggestion into proof. Among the most important and interesting of such experiments we must place those of Professor Correns with two species of the Bryony plant. In the species known as Bryonia dioica female flowers are found on one plant and male flowers on another; the two sexes are borne on different individuals, so that any particular plant is either male or female, and not hermaphrodite like the majority of plants. The other species, named Bryonia alba, has the sexes borne on different flowers but on the same plant. Each individual plant is therefore hermaphrodite, bearing both male flowers and. female flowers. When the flowers of a J 130 THE MENDEL JOURNAL female B. dioica are crossed with those of a male B. dioica, the offspring consist of a mixture of male and female plants. The most interesting crosses, however, are the reciprocals between the hermaphrodite plant of B. alba and the two unisexual plants of B. dioica. When the flowers of the female plant of B. dioica are crossed with the male flowers of B. alba, all the in- dividuals in the offspring are female plants.* But the reciprocal cross, strangely enough, gives a very different result. For, when the female flowers of B. alba are pollinated with pollen from the flowers of a male B. dioica, the offspring consist of male and female plants in approximately equal numbers. Correns endeavoured to interpret these results in the following way. Let us take first the cross of male B. dioica with female B. dioica. The result, as we have seen, Is a mixture of males and females in equal numbers. We will for the moment assume, as Correns did, that maleness is dominant ; that the male plants are Mendelian hybrids and therefore carry two kinds of pollen-cells in approximately equal numbers, one half bearing the character of maleness and the other half that of femaleness; and that the female plant, being recessive, must be pure with regard to femaleness, and all its egg-cells will therefore carry femaleness alone. It is clear that a cross of the *A few exceptions—2 males with 589 females——are said to occur, and a few of the females bear occasional male flowers. MENDELISM AND SEX 131 nature which we are now considering will resolve itself into the simple Mendelian one of DR by Rk.* The DR here represents the male plant and the RR the female plant. The symbol D in this case stands for maleness, which is dominant, and R for femaleness, which is recessive. Since the female plant bears only “ female” egg-cells and the male plant bears both “male” and “female” pollen- cells in equal numbers, it must happen in the random meetings resulting from pollination that one-half of the “female” egg-cells will be fertilised with “* male” pollen-cells which will give us D# offspring, and one-half with “female” pollen-cells which will give us RR offspring. And, since maleness is dominant, then the individuals which result from a fertilisation of R egg-cells by D pollen-cells will be hybrids and will be visibly male. Thus there will be produced equal numbers of hybrid males (D F’s) and of pure females (R #’s). So far, then, the Mendelian interpretation is in accordance with the experimental facts, and it gives them an in- telligible unity. Let us pass next to consider the crosses between the two different species. As we have already seen, they resolve themselves into crosses of the two following kinds :— NATURE OF CROSS. Bryonta Droica.