The Genetics of Short Fourth Toes in Humans in India

THE GENETICS OF SHORT FOURTH TOES IN HUMANS IN INDIA

By AjIT KISOK I~.AY Ge~zeli(;s and BiomelO~ Laboralor)~, 6'ove,'m~e~l.t qfi Orissa, Bhzl.baneswar-3, Orissa, India.

Kay and Haldane (1965) have published a preliminary account of the abnormal shortening of tim fourth toe ill humans, occasionally accompanied by other skeletal deFccts. It had beer~. previously reported fl'om Japan (Katsura and Hayakawa, 1938; Ogawa, 1953; and ochers) where, though l~ot rare, it was not as common, as in Ii~dia. It has also been reported fl'om Yuc~'lta.n (Steggcrda, 1942), but is very rare it" not absent ill Europe and. North America. While screening males ibr colour-vision in villages in Orissa, of 2500 mini examined, th.is abnormality was observed in three of them. Since then I have looked for it ill other states. As most people in Indian villages do not wear shoes or sandals, the condition is easily observed. The con.dillon was found to be genetical, and several pedigrees have been collected. I do not think this abnormality is due to malnutrition or other environmental e.ffects. The di,fferences in the occurrence of the condition in India may be either due to gene Kequellcy differences in the main gene (or genes) or in modifying genes. The condition was reported in Ja.pan by I=iiroki (1911). Hie found two cases among 4804- out-patients in the orthopaedic clinic of Tokyo University. Further cases have been reported by Katsura and Hayakawa (1938), Ogawa (1953), and Sugiura (1961). .Bell (1951) eolatains two pedigrees. The first was reporeed by Steggerda (1942) ill Yuektan, where he found bilateral shortening of the fourth metatarsals in two sibs and their maternal uncle. The second, reported by Birkenfeld (1928), contained three M-reeled individuals, two sibs and their mother. Gates (1946) cited a few pedigrees of fourth short metacarpals, the condition being an irregular autosomal dominant, and occasionally accompanied by short fourth toes. In the pedigrees reported in this paper, the shortening of the fourth metatarsal is associated with other abnormalities in only 8 % of cases. In India. several superstitiolas are associated with. this defect, some believing it to be a good sign and some a bad one. Most of these superstitions refer to the period ol'preg- nancy, i.e. either because the mother did not obey certain food or other restrictions, or because she wellt out of the house on the night o[' a lunar eclipse. Anoth.er belief is that if a woman has several still births, and if she cuts o~' the fburth, toe of the last still- born child, sh.e will have a live birth and that this child will have short fourth toes. Others consider a female child with short fourth, toes as Lalcshmi or goddess of wealth and of the lmme. Some people also prefer to marry their daughters into families in which AjIT KISOR RAY 9c~

this abrmrmality is known to occur. These beliefs regarding children indicate that thi conditiol~ is commonly known to be congenital.

DATA GOLLEGTION

The data were collected most:ly in the states of"West Bengal and Orissa. OF the 81 propositi, 36 were born in West Bengal, 28 in Orissa, 13 ill other states of India and 1 ia East Pakistan. The classification of the propositi accovdi~zg to endogamous group is as follows.

West Bengal : Brahmin--2; Kayastha---8; Kora tribe--1 ; Kshatriya.--1 Mahisya--3 ; Muslim--ll ; Pramanik--I ; Tanfi--1, Total--2~ Orissa. : Brahmin--2; Gaura--1; Khandayat 4; Muslim---3; Pano--1 Tell--4. Total--15. East Pakistan : Brahmin--2; Kayastha--3; Namasudra--5. Total--10. Other states : Goanese--I ; Koya Dora tribe---2 ; I-Iadi--1 ; iL\/lalayali (Kerala)- 1 ; Telegu (Andhra)--4. Total--9.

For the other 26 propositi, such data could not be collected., including 23 cases whic could not be followed up. The above grouping' is according to caste, language ( religion, Most of tffese endogamous groups are further divid.ed into endogamous sul groups which I ,have not given, In villages, where most people do not wear any shoes or sandals, collection ot' da, was velatively easy, whereas in a town, even if" people wear light and open sandals, tl partial view afforded is not enough for a diagnosis, l~/Iost pedigrees recorded in tl~ paper were collected by talking to villagers about the abnormality.

PEDml~EES (S~E AI'PENDIX)

A. A/iuslims--Cut tack district (Orissa). As there were ve W ['ew Muslim conversions in Orissa, t-he Muslims are an endogamo group forming a small minority of the population. The pedigree contains 7 affect~ males and no affected ['emales in four generations. 3 o1~ them are affected bilateral and 4 unilaterally. The pedigree includes two first cousin marriages. The proposit (III.12) had 12 children fi'om two wives; two sons were affected. Two of his brothe were affected, and his ['ather and grand['ather were also reported to be affected. .~ the 7 a~ffccted individuals, all the 5 still alive were personally examined.

B. d14uslims--Cutta.ck district (Orissa). This pedigree is related to pedigree A and the members belong to an adjoining villa in the same district. Four males, 2 symnaetrically and 2 unilaterally and two femal, olle symmetrically and the other unilaterally, were affected. Two of" the cases we examined, 4 reported, including 2 dead. 100 Shorl fourlh loes ia humalz,~" c:-t-c',o. Tal~ti--C',aleutta. The two propositi were indepel:den.tly discovered, and. are :'elated by marriage. They are not known to be otherwise related. The caste is subdivided it:to smaller sectiol:s, generally territorial. The pedigrees include 4 male and 4 fen:ale a.l:tEcted individuals, all but one being affected bilaterally. Only fen:ales have transmitted the condition. l~ive cases were exam.ined and three reported.

D l-lDo. It~wasthas--C, aleutta.. The members of this pedigree were originally fi'om Dacca and Barisal in East Paldst:an, but since the partitioning of Bengal in 1947, tl:ey have lived it~ Calcutta. They are connected by a marriage. They beloag to the Ba~zga]a sub-caste of the Kayasthas. The Ba~zgq]as belong mostly to East Bengal. This pedigree contail:s the largest ram:bet of affected individuals 9 females and 2 :::ales i1:5 generations. Five females were affected uMlateralO,, all o1: the right foot. Six il:dividuals were examined, and. 5 were reported, including 3 dead.

13. Brakmhls--Puri district (Orissa). A former King of Puri is said to have induced several Brahmins ofJajpur (Cuttaek district), who had migrated fi'om Kanauj, to settle round the l:ow famous temple of Puri and carry out the religious ceremonies connected with the idol Jagannath. The Brahmins who came to Puri for this purpose are called the Dakshi~lal,ya Sre~zi (or Southern class) and those who stayed behind at Jajpur are the Ultara Stem (Northern class). These sul>castes are.further sub-divided il:to smaller endogamous groups according to the functions they perform in the tetrq?le rituals. The pedigree contains 4 generations with two males and one female affected.

Y. Telis--Cuttaek district (Orissa). The caste is subdivided into smaller el:dogamous groups. Two males and one female were affected, all o1: the right foot only. One was examined and the others dead.

G'. tgayasthas--Caleutta. All the affected members of this pedigree except the propositus are living in East Pakistan. The proposims migrated to C,aleutta after 194.7. In all, two males and two females were a~'eeted, all unilaterally.

:H. Brahmbzs--C, alcutta. This family I~r initially to Faridabad (East Pakistan), but migrated to C,alcutta after 1947. Two males and. 9 females were affected, all bilaterally. AJIT KISOR RAY 101

L ll/[akilwar~lf--Cuttack (Orissa). The 21/-[akilzoarus are leather dressers and. cobblers fl'Oll?, Andhra State. This famil is ~mw settled in Cuttaek. The pedigree COlltains two apparently unrelated sol tree of this condition. Ill all, there are two males ali'ected, bilaterally and one unilaceral[3 mid one female affected unilaterally. In addition, one female in the pedigree wa reported to have short fbul'th fingers on both hands. j. Telis--C, ut:tack (Orissa). Two males and fbur females are affected bilaterally and one male unilaterally. On case was examined alld six were reported, in.eluding 4 dead.

K. Chrislians~(Kerala). The pedigree was collected at Poona, and. consists of two all%cted sisters, one o['whol was reported.

L. Telis--C, uttack (Orissa). Two males al~.d olle female were a.fl.'ected, all bilateral.ly. One was examined, tl other two being d.ead.

"IVL Kayaslhas--Calcutta. The two affected females had Iollg illdex ~n.gel's, attd one una.li'ected male had pol dactyIy on all four limbs.

N. k4llslhns--Puri (Orissa). Two males are afLEcted, bilaterally. Both were exam.ined.

O. Panos--Pm'i district (Orissa). The Pano caste are considered to be remnants of an aboriginal tl'ibe who beca~ Hindus. They arc considered a very low caste. In different parts of the state, th are occupied in d.i~'erellt ways, e.g'. weavers, baskee makers, night watchmen, musiciaJ aud palallquin bearers. Each of these units is strictly endogamous. Three att'ect males and. one aft'ectdd female belonged to three generations. Two of the males shox.~ the condition bilaterally. Three of" the affected ir~d.ivid.uals were examined al~d c was dead.

P. Gauras--Puri district (Orissa). The Ga,tras are a pastoral caste of Orissa and one sub-divided into sevel'a[ sub-casl One of" the earliest settlemelltS was neat' M.athtu'a (U.P.) and they claim descm~.t fi', Krishna. Three males in. three generations showed tile ablaormality, one of th un, ilatel'ally. Two cases were examined and m~e reported. 102 Short foz~rth toes i~z hz~ma~zs

Q. Telis--Puri district (Orissa). Fore' males were affected bilaterally and one female unila.terally. Four affected individuals were examined and one was dead.

~. ll/Iahislo,as--Burdwan district (West Bengal). Two females and one male, all of one sibship, were bilaterally affected, and were all examined.

S. Brahmi~s--Burdwan clistrict (West Bengal). The two a.ffected females, mother and daughter, also showed syndactyly of the second and third toes on both feet. Both were examined.

T. tCcoJaslhas---Bm'dwan district (West Bengal). Two sisters were bilaterally affected, one of whom was examined.

U. l]~uslims--Burdwan clistrict (West Bengal). Three females were affected, two of them bilaterally, Two males were aP2ected bilaterally. .One of the latter also had a short third toe on the right foot. Three of them were examinecl and two were dead.

V. l~/f~lslims Burclwan district (~fest Bengal). A male and a female were affected, both on the left foot only. The affected female also had short t-erminal phalanges of the thumbs. The male was examined, the female reported.

W. l]/fzlslims--Burch.van (West Bengal). Two males and one female were affectecl, all of them on the right foot only. One nmle was examinec[ anc[ the others were dead.

X. Namasu&as--Burclwan (West Bengal). The family belonged initially to Varidabad (East Pakistan). Two males and two females were affected, all bilaterally. An unaffected male had polydactyly of both hands and another had syndactyly of the third and fourth :fingers of the right hand. Only one male was examined.

Y. --Burdwan district (West Bengal). Three females were affected, all on the left foot only of whom only one was examined.

Z. Koras--Burch,van district (West Bengal). The t~oras are a tribe who inhabit mainly Chota Nagpur and the Santal Paraganas. The th.mily has lived in Bm'dwan for three generations. Two males and a female were bilaterally a~'ected. The two males were examined. AjIT KISOR RAy 10~

AA. el#lMims'--24 Pargan.as district (West Bengal.). Three mates and 2 females were affected, all bilaterally. One al}'ected female ha( sliort ttmmbs, and an una~Ected male had short great: toes. Two affected individual were examin.cd, and three were reported, including one dead:

AB, Ii'r alcutta. This family belonged to M~idnapur district (West Bengal), and are now settled i~ Calcutta. Seven males and. one female were affected. One male was ali'ccted ou.l' on the /.eft foot:. Two affected individuals were examined and six were reported including three dead.

AC. Hadis---C,alcutta. The Hadis are an endogamous caste of giha.r. The family lives in Calcutta. Thre males and two females were affected, all bilaterally. On.e affected, male and on aiTccted female had short thumbs. An. un.aiTected female had a short left- thumb. Tw affected individuals were examined, and 3 dead..

AD. t{'wasfflas--Calcutta. A male and 1l females were affected, all bilaterally. The pedigree contains tw genetically in.dependent sources of the abnormality. Three a;ffected individuals wer examined, and 9 were reported, in.eluding one d.ead.

AE. Ifnj,aslhas--C, alcutta.

Three individuals were affected, all in the right foot alone. Only one case wf. examined, one of the others being dead.

AF. tgc~,asthas--C,alcutta. The f~tmily belongs initially to Faridabad (East Pakistan). Two sisters were unil~ terally affected. On.ly one of them was examined.

AG. Kdlalri2,as--Berhampore (West Bengal). Two males and 3 females were a~l:Ihcted. Only one female has both fourth to short. Two a~ected individuals were examined, and 3 were reported, including m dead.

AH. .A%oJast/las--Berllamlgore (West Bengal}. Th.e pedigree includes one male affected on the right tbot only and a female wil both fourth toes short. Only the former was examined.

AI. ,A@~ncar~dras--Bcrhampore (West Bengal). This pedigree also includes one male affected on the right foot only and. a. fema affected, bilaterally. Only the tbrmer was examined. 104 Short fourth toes in humans

AJ. Bikari--Calcutta. Two males and a female were all affected, oll the left fbot alone. Only one male was examined.

AK. --Calcutta. The family was originally fi'om /Vl.idnapur, and are now settled in Calcutta. Two ~nales were affected bilatc~'ally, one of whom was examined.

AL. --Oaya (Bihar). The pedigree was recorded at C,alcutta, and included three males and two females affected, all bilaterally. One male only was examined.

]) ISCIUSSION

An examinatim~ of the pedigrees A to AL leaves little doubt that the condition depends in part oll an autosomal gene whose pcnetrance in heterozygotes is not very high, i.e. it behaves as an irregular autosomal dominant. Of the 61 pedigrees, 38 contain at least two affected persons. ~[n addition I have seen 23 cases where I could not get any information about relatives. Four of the pedigrees contain two independent sources of the gene. In all, I have observed in India 117 persons who had short fourth toes on one o1' both of their feet. Besides these, 51 living and 33 dead persons were reported to me to have had this condition, making a total o[ 206 cases. In 130 cases, both feet were a.ffected, in 40 the right foot alone and in the other 36 the left foot almm. 104 men and 102 women were affected. Ikadiograms of 15 of the affected individuals disclosed short [om'th :metatarsals, which were about as long as the l~fth. In most cases, the shafts of the metatarsals were shorter and broader than normal. Fourth short toes were commonly accmnpanied by other skeletal abnormalities. The ones recorded in the pedigrees win'e: 4. individuals with short terminal phalanges bilaterally; 3 with mmsually long fingers on both hands; 2 with s3qxdactyly of second and third toes on both feet; 2 with short third toes on one foot. Some members with normal fourth toes also had skeletal abnormalities as follows: short great toes; short terminal phalanx unilaterally; short fourth fingers; polydactyly on hands and :feet; club feet; short third toes; syndactyly of third and fourth fingers of right hand. The following associated abnormalities were seen among affected individuals who are not included in the pedigrees : patella absmlt on the sa:me side as short fourth toe in addition to dental abnormalities; ~t much reduced patelh on the same side as the short fom'th toe and polydactyly on a hand; short third toes; long little :finger. 1~ suspect that xnost of these abnormalities are manifhstations of the Inain gene, which is affected by d.ifferent modifying genes in different pal'ts of lipidic. In Japan, the condition is fi'equently accmnpanied by short fourth and :fifth cmetacarpals, the ;fifth mesoi~halanx or the first AjIT KISOR RAY 105

telephalanx of the hand. Bell (1951, Plate V, 27, 28) records that short metatarsal: and metacarpals were found in an adult achondroplasiac. The fi-equeacy of the condition varies greatly in di~'erene parts of India mad amont ditTerent endogamous groups. I a 11 villages in ~a,rest Bengal in the districts of Burdwan J3erhampore and 24-Parganas, the population was surveyed tkirly exhatislively. probably naissed some slay middle aged women, especially Muslims who are very con servative. A.s 33 affected men and 24 affected wome~ were discovered, this bias i probably s:mall. Because of immigrations fi'om East Pakistan in recent years, th populations of these villages are uncertain. I~ a population of 7400, I observed 4 living cases, giving a fi'equcncy of about 0.66 per cent. I:['we exclude fi'om this nl_tnlbel 3 aboriginals and. East Pakistalfi relhgees and their children numbering 11, lhe fi'e cluency is insignificantly reduced to 0.60 per cent. In villages in Orissa, 3 cases wer found, among 2500 persons examined and among the Koya Dora tribe in Andhra 2 persons out of about 1000 to 1300 were a,ffected. This gives a fi'equency of abot 0"13 per cent. As the gene has very low penetrance, the gene fi'equency is given by lh figm'e divided by twice the penelranee which gives the value of 0"18 per cent. T17 fi.eque:ncy may range widely in India. Tanaka (1962) states that the abnormality is ve W rare in Japan, He found abnormals among 3115 school girls exalnilmd. This gives a fi'equency of 0"1%. Iq found no cases in a comparable sample of boys. On present evidence, there is not" a excess of females among the Indian cases. I-Iowever, as men's feet are more ofte seen than women's, there is a large excess of males among propositi. Tanaka's obsevv~ tions also show that the condition can be manifested in gMs of school-going age. t India I have not observed the condition in anyone below the age of five, but there ai superstitions referring to its presence at birth.

Table 1 (a). Abnormals among first degree relaUves of propositi

,m e

t/l

c? 36 10 5 7+7 ,1 28 1 4. Total 64 11 9 13 98 10 79 11 160 27 [37 Table 1 (b). Sexes of abnormal first degree relatives Relatives

28 23 51 Propositus 7 23 30 35 46 81 106 Short fozoth toes in hzmzans

From my data, one eaa calculate the penetranee of the gene. In the case o('a l~r~ fully penetrant rare gmle, half the parents, half the siblings and. half' the parents o{'{a~ a.iTected individual should be affected on the average. It'we observe that less than liaie the relatives of' the above kinds are affected, the gene is pr01?ably partially penetrafit Iffowever, since I have deliberately looked for relatives ofpropositi, the estima.te obtail~ed from th.ese pedigrees will be too high. In pedigrees with only one affected memberl dm penetranee estimate is zero. This will provide too low a.n estimate. Tables 1 to4 summarise data ill order to calculate penetrance estimates. ]if the pelmtrmaee of ihe gene is p, the proportion of :first: degree relatives (parents, childrert al~,d sibs) of' al-t affected person that we expect to also show the character is 1@ Similarly, we expec~ i]:P of grandparents, grandchildren, m~cles and auras, nephews aad nieces, aad l',ai~ sibs ot` affected persons to be aflket'ed. I have not ilieluded ia my pedigrees, dead relatives of living iadividuals more remote thal~ parents. All secmad degree relative s of' propositi are not therefore inelnded ill the pedigrees and I use the term leeent ab~mrmals'.

Table 2. Pe~letranceeslimale Ji'om firsl degree relatives of propositi Abnormal Total % p Parents 20 128 15.6 0.3125 Clhildren 23 177 13.0 @2599 -2691 Siblings 38 297 12.8 0.2559 81 602 13'4.5 The proportion of first degree relatives that are affected is 13'45Yo. The estimate o~ peaetranee is 0'2691 (see TaMe 2). The penetrance estimate [br different types of first degree relatives varies from 0.2559 for siblings to 0.3125 for parents. The estimates from the second degree relatives are less reliable. I have only included i1~ the ealeula: lions those grandparents for whom I could obtain infomnation, although it could be argued thai: if one parent is affected, the pm:ents of the other parent can be assumed to be normal. The penetrance estimate fi'om all second degree relatives is 0'2301 (seeTable 3); the estimates for differen.t types of second degree relatives vary from 0.1404 for graudpareats to 0'8000 for halt`-sil3s.

Table 3. Ahsormals amo~g second degree relatives Grand Grand Uncles & Nephews & Half brothers parents children Aunts Nieces & sisters

m m m m m

< 4. 114. 1 10 16 195 9 215 1 5 o abnormal 3.51 10.00 8.21 4.19 20'00 Penetranee estimate 0.1404 0.4.000 0'3282 0' 1674 0'8000 Combined peaetranee cstlmate=0-2B01 AJIT KISOR RAY 107

Lf: we take all sibships con.tainin.g an abnormal whose parents are included itz the pedigiiee , 4.7 of the 117 sibships have one abnormal parent. The estimate ofpenetrance 0~taincd is, theretbre 47/117 or 0'4.027. So we can say with some confidence that the vakm o[" penetrance is between 23 % and 40%,

Table 4(a), Sex o/' aff'ected .parenl,-off27)rin,g pair

Off:spring

30 16 4.6 Parents ? 15 20 35 4,5 36 81

Table 4(b), (Pedigrees A, AB, AD omitted)

18 15 33 14 11 25 32 26 58

The numbers of transmission Of the abnormality are given in Table 4, The value c

X t for Table 4(a) is 4'02 (P>'05). This departure fi'om independence, i.e., ~fidency to transmit the trait to the same sex more often than to the other is mainl due to the three pedigrees A, AB anti AD. If these are removed (Table 4(b)) ,~ =0'l~ ~eis possible that a di.fferent locus is involved in these three pedigrees, but tbr the put poses of this paper I assume all pedigrees to involve the same locus.

Table 5, Laterality in Parents and offsprings OKsprlng Parent RL R L O Total p lkL 46 5 5 189 245 .4571

R 6 11 0 71 88 ,3864 ,3676

L 1 2 5 40 48 ,3333

I next consider the inheritance of asymmetry and laterality. Considering paren offspring pairs, Tables 5 to 7 summarise the data. The penetrance among childr( Of bilaterally affected parents is 0.4571, that among cl~ildren of asymmetrically affectc parents is 0.3676. The difference is not significant (P=0'30). From Table 6, X21 22'4:, which is very highly significant. Therefore symmetrically affected parents ha' a greater chance of producing symmetrically affected children and vice versa. t08 Shot} foz~rth lots bz hu,ma~s

Table 6. Symmelr), at~d asymmel O, i1~ parelzl,s atzd offsflvings O f[i~prlng SYM ASY SYM 46 l0 56 Parenl: ASY 18 25

53 28 81

Table 7. dsymmelr), ilz parelzls alzd o~;71rilzgs

O~tSprlng

R L

R. 11 0 Parent L -2 5

I have naturally looked for a homozygote. One might be expected in pedigrees A, C, D and AD, due to inbreeding in A and because there are two independent sources: of the gel~e in the other 'three. In pedigree A, there were two mmTiages in each of which one parent had an affected tSther, and the or'her an affected grandfather: Therefore, the probability that both parents in either marriage were l{eterozygous is s.:• These two marriages together produce 5 children; therefore the probability that one of these children is homozygous is 5 y, .z-~,• ,~ Similarly, computing this probability for all the relevant pedigrees gives us the probability of linding a homozygote as ~,~-,"a- To have a high probability of lincling at least one homozygote, we should need about live times as many pedigrees. Of course, the homozygote could be lethal at an early stage and tmlikely to be reported. It could be lethal soon before or after birth, gravely deformed, or like the heterozygote. The last alternative seems most likely.

SUMMARY

61 pedigrees of short fourth metatarsi have beenrecorded fl'om India. 38 pedigrees containing more than one person and 23 null pedigrees. In 130 cases the anomaly was present bilaterally and in 76 unilaterally, The largest pedigree includes 12 affected melnbers and the longest: has affected members of tive generations, 117 persons were personally observed besides 51 living and 38 dead short toed members; The data are consistent with the gent being an irregular autosomal dominant. About 8 per cent of affected persons have some other associated anomaly. The true value of penetrance is very probably between 23% and 40% and the penetrance tMls o~ with 'distance' fl'om the case. There is good. reason tO believe the inheritance of sex ineiden.ce in some pedigrees and for laterality. A.IIT KIsoa RaY 109

AC;KNOWLEDGE/vIENTS

I1~ the preparation of this paper I have profited greatly Ii'om the Idnd suggestions arid commen.ts of Mr. S. D. Jayakar. I am indebted to the late Professor J. B. S. I-Ialda.ne, F.R..S., alld Dr. Helen Spurway for encouragement and stimulus i~l my work. I wish to take this opportunity in than.king all those who assisted me in my field work, ia particular, Mrs. Anima Kay and Shri Kripa Sindhu Sahu, who provided me with t:he names of ehree propositi.

RE FEI?,-ENCES B~LL, .J. (195 I). The ~'ea.~wg'oj'I-inman Inheritance, Vol. V, Part I. CamlJridge University Press: London. BmKI~NVELIJ, (1928). Uber die Erbllchkeit der 13rachyphalangie. Arch. Klin. Chit., 151, 61 l-a 1. GATES, R. R. (1946). I-Iuman Gomllvs, 17oL 1. The Macmillan Company: New York. H~KOKI, B. (1911). Statistics of congenital mal[brmatlons. Totg,o, Igatckai-zasshi, 25. KATSUaA, S. aND HAYaZCAWA,S. (1938). Short tburth metatarsals. .hCisshln-Igaku, 27, 1499 m~d 1651. OoA~VA, S. (1953). Family study of the'short fourth metatarsals. J. Japan. Or&op. Surg. Sot., 27, 83. RaY, A. K. aND HaLDaNI~, J. B. S. (1965). The genetics o[' common Indian digital abnormality. Proc. Nal. Acad. Sci. U.S.A., 53, 1050-1053. STEGClERDA, 1~4. (1942). J, IZfered., 33, 233-234. Summ~A, (1961). On the skeletal variants in hands and feet of Japanese children. J. Jap. OrlhoD. Assoc. 35, 925-6. TAN:tiC.A,K. (1962), Differences bct\vcen Ciaucasians and Japanese in the incidence of certain abnormalities. !it The 6'enelics of~l/Iigran! and Isolale P@ulallons. The Williams and Wilkins Company: Nm~ York. 110 Short fou'ri/z ~oes in humans

no cht7

B AJIT KIso~ RAY 11 1 112 Slzort J'oz~rUz loes 7n, kzu~al~s

d 4-,~ -Fd d d +, +d ~, +~ +

shorl` fourth toe ossociabecI :::: wil-h 3rd shof? koe of It, fool-

chil. AIIT KIso~ I~y 11

8 i i4. S/zort fou~l/z toes i~z /zzl~na'~zs

~) short fourth fingers

J AJIT KlSOR I~AY 1 1

shorl- 3rd toe on.boil7 fee/ ].16 Short Jburtlz toes i~7, human,s

d I I

4•)-po/ydacty/e on bo?h hands,feet ~i~ short fourth toe associated with long index fingers

d AjIT K!soR RAY 1]

short fourth toe assodoted ~/ith long fndex finqers r ~econd toes

/ 1t8 Short fourth toes ia humans

R AJIT Kisolz RaY 1

short fourth toe associc~ted wifh syndactyle : 2nd~r?rd toes of both feet

T 120 Skort fourth toes i~z kumans

d d

@snoft fourth toe osSOCl'gted w/th short 3rd toe of r#. foot

@ short fourth foe assoc/atecl with sllorf thumbs AJIT KIsor~ RAY 12

14/

@ po/ydacty/e on. bdth hands 0 syndacty/e, 3Fdf4th fingers of r~. hand 122 Sl~orl fourll~ foes hz huma'lzs

% AjIT I~ISOR RAY 12~

~~~ ~

5,~,SSS

| short greot toes @ short fourth toe associated with short thumbs

'4 124 Short fourth toes in humans

,4C

0 shark thumb of It: hand @ shark fouckh toe associal wffh sl~ort thumbs

AD Ajn ~ K~soR RAY 12E

AF ]26 S/~orl fourth toes hz h,uma~zs

A/-/ AjIT KISOR RAY 12;

~f Gj~~

Ad- 128 Short fourth loes in humans

,4/4

AL AjIT KlSOR RAY 1~

b 1.30 Short fourth toes bz humam"

~(~ ~ polydacty/eon It..hand adjoininq t't~umb

d AjIT KlsoR RAY 13

O syndackyle fourth t fifth toes on both feet i 32 Short fourth loes i~ hztmans

h Ajrr KmoR RAY 13~

d--9 134 Short Jburth toes in humans

k A IIT KIsoz I~AY 1,3~

n ~d 136 Skort fourt/t, ~oes i~z kuma~s

P AjIT KIsol~ RAY 13

? F

10 138 Short fou,'th toes in hu,nans

r short fouFth toe of It. foot associated with short third toe of rt, foot

t AJIT KIsoI~ 1~.u 1 t/

no cMI.

V 140 Short fourth toes i~z humans

Vv'

d? I?Or'ma l short fourth toe on both feet

short fourth toe on Rt. foot

Short fouFth toe on Lt. foot

exam~ned

d dead

other assoc/ated abnormal/t/es

reported /n the pedigrees