THE INHERITANCE OF MEDULLATION IN WOOL
SING SZE YOUNG
A THESIS
submitted to
OREGON ST&TE COLLEGE
in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE
June 1949 APPROVED:
Professof of Aninia1\tisbandry In Charge of Major j
Head of Department of Animal
Chairman of School Graduate Committee
Dean of Graduate School ACKNOWLEDGEWNT
The writer desires to acknowledge his thanks to
Dr. Ralph Bogart of the Department of Animal Husbandry for his precious guidance in preparing this paper.
His sincere thanks are extended to Professor
Oran M. Nelson also of this department in giving much helpful instruction in the collecting of material and laboratory work.
The writer also wishes to express his gratitude to Dr. James N. Shaw of the Veterinary Medicine Depart- ment for granting the facilities for making pictures of all the samples. The saine gratitude is also extended to Dr. 3erome C. R. Li of the Mathematical Department for his advice on the statistical work. CONTENTS
I. INTRODUCTION AND LITERATURE i
II. MaTERIAL AND THOD 4
III. RESULTS 6
1. Statistical Analysis
IV. DISCUSSION 17
V. SU1VIM&RY AND CONCLUS ION 25
VI BIBLIOGRAPHY 27
VII. APPENDIX 29 - 1
T1E INHERITANCE OF DULLATION IN WOOL
I. INTRODUCTION AND LITERATURE
Medullatlon in wool has been revealed in early days (15, p.26). There are two kinds of wool fibers that posses medulles: the true, short kemp and the inedullated hairy fibers. Kemps are fibers that grow for a definite, limited time, and. are shed sooner or later after corn- pleting their growth. Sometimes hairy fibers that are not shed, but are of intermediate growth, are produced in follicles which have previously given rise to one or more kenips. Med.ullated hairy fibers of the non-kemp group are those of interminable growth which are the first occupants of their follicles. The non-kemp niedullated hairy fibers may grow in one of two ways: (1) as the successor of kemps or (2) directly from the original follicles. The causes of medulla formation are still not exactly clear. Several hypotheses have been launched to explain its occurrence, among these are: nutri- tional, shearing and. hereditary. 2
Waters and Dry (3, p.2?4) found freedom from medulla in the layer of the fleece of certain animals which was grown very slowly in winter, while there were many medullas in the parts grown more rapidly before and afterwards. According to this hypothesis medulla forma- tion is attributed to the vigor of the fiber root, which in the above case was thought to be putting out material faster than it can keratinize this fiber stuff to form the substance of pure wool.
Rudal]. (19, p.28) in his experiment on the effect of shetring reports: "of the differentially shorn anima1 in each ease more hairiness appeared In the shorn side than on the unshorn side". The cause, according to him, is due to the exposure after shearing and. Is not due to the mechanical effect of shearing.
The hereditary cause of medullated fiber has been considered early by Bryant (1, p.184) in her work with the Scotlsh Mountain Blaokface sheep. That the hereditary factor Is the cause of medullated. fibers is partially vers fled by Wilson (20, p.142). Sections through the epider- mis of the thigh region were obtained from several Romney embryos taken late in gestation. The presence of niedul- lated fiber were clearly shown, indicating that the medulla Is formed before nutritional or enviroruxiìental fac- tors have had opportunity to become causative. 3
Duerden (7, p.273) in discussIng the origin ot
hairiness attributes the presence of hairy fibers to the
railure of the oriina1 outer protective coat of hair en-
tirely to disappear in the process of evolution. He re-
ca11s the ract that in their natural state nearly all
mammals have two types of hair; an outer coat of long,
coarse fibers an an under coat o' short fibers.
This two-coat type of fleece is still possessed by most
of the rather primitive Asiatic sheep.
Duerden evidently considers the problem of elimi-
nation of the medullated. fibers as lying outside the field
of nutrition, for he states: "cornplete elimination should
be possible by continued selection in breeding".
Dry (0,6, p.332-338) later developed the idea
that wool fiber can be classified into five different fiber-type arrays in the evolutionary meaning. He thinks that hairiness or the formation of the medulla is one of the outward manifestations of forces shaping the structure and development of fleece fibers. These forces, which he called pro-natal check, are responsible for the evolution of the fleece from mixed or two-coat covering to one-coat covering.
Based on the fiber-type arrays, Galpin (12, p.l2 has been able to study the britch-poll gradient of the Romn.ey lamb fleece. She reports: "some medullation oc- curred in all of the Roxnney coats examined, but there was marked variation in the percentage found in different coats. A higher percentage of niedullation was found in coats with slightly checked arrays and a lower percentage in those coats where the arrays were more checked. In the Southdown breed, medullation was negligible; it IB more common in the Ryeland breed". In this country not much work has been done on this subjeot because medullatíon Is almost absent in the fine wool breeds such as Merino and Raìnboulllet. But due to the growing number of the Romney sheep in the Willamette Valley and the surprising number of medullated. fibers in their fleeces, it is of much interest to study this character. The purpose of this study is to determine if niedullation is Inherited. Also, the mode of inheritance of this characteristic will be studied in so far as the data available will permit. Besides, the methods of elimination of this defect in wool are discussed.
II. MATERIAL AND LThTHOD
sour raras were chosen representing non-niedul- lated., medium medullated. and heavily medullated types 5 respectively. Ram A-86, a Roniriey, and. ram H-279, a Hanip- shire, both are free from medullation. Ram C, a corder Lelcester, has a inedullatlon of one per cent on shoulder and 18 per cent on the britch and was considered as medium niedullated. Ham D, a Cheviot, has a Inedullation of 22 per cent on the shoulder and 100 per cent on the britch and was considered as heavily niedullated. Seventy-eight cross breed ewes o1 Lincoln i Ramboulliet, Romney x (Lincoln x Rambouillet) were bred to the above rams. Half of the ewes were free from medulla- tion and the other half have a medullation ranging from one per cent to 50 per cent (only one ewe) are considered as medium medullated. Wool samples were taken both from shoulder and britch from those offspring born froni the above breeding as lambs. The ewes when sampled had mature fleeces. All ewes and lambs were kept under the same environment. The fiber samples were cross-sectioned by means on 01' the Hardy's device (ls, p.385) and were projected a printing paper at the magnitude of 250 times the actual size. .rom the magnified picture, 100 fibers were counted out as a representative and were classified by types, that is, non-medullated and medullated fibers. Either pin-head medullation or large niedullation were counted as medullated fibers. There were not many Icemps in these samples and therefore, they were disre- garded.
III. RESULTS
The percentage of medullated fibers was deter- mined after the samples were cross-sectioned and pictures made of each. Figure 1 and figure 2 are presented as repre- sentative of the two extremely different tTpes of niedul- lation; one is loo per cent medullated and the other is free from niedullation. Representative pictures of britch of the sire and dam and resulting offspring for the six dit- ferent kinds of matings are shown in figures 3, 4, 5, 6,
? and 8. Non-medullated rams mated to non-medullated ewes produced lambs averaging 5 per cent nied.ullation. (Figure 3) . Non-mnedullated rams mated to ewes with medium medullation produced lambs averaging 14.6 per cent medullation. (Figure 4). Medium niedullated rams mated to non-medullated ewes produced lambs averaging 21.7 per cent medullat ion. (Figure 5). Medium medul- lated rams mated to ewes with medium medullation pro- duced lambs averaging 58. per cent medullation. (Figure 6). Heavily medu1late rams mated to non-medullated 7
Figure 1. Cross section of 100 % inedullated wool (250 X).
Figure 2. Cross section of non- medullated wool (250 X). roi LJ
Ewe
Non-medull ated Non-medullated
Lamb
5% medullation
Figure 3. Non-medullated ram x non-meduliated ewe
Ewe
Non-medullated Medium medullated
Lamb
14.6% medullation
Figure 4. Non-medullated ram x Medium medullated ewe R am Ewe
Medium-meciullated Non-medul
Lamb
21.7% medullation
Figure 5. Medium-medullated ram x Non-medullated ewe
Ram Ewe
Medium-medullated Medium-medullated
58.3% medullation
Figure 6.. Medium-medullated ram x medium-medullated ewe. Ii 'k%TvLJ r X
Non-medullated
76% medullation
Figure 7. Heavily-medullated ram x Non-medullated ewe
Ewe i$ w
X
Medium- medullated
Lamb
Medullation
Figure 8. Reavily-medullated ram x Medium-medullated ewe TABLE I. MEDULLATION IN THE WOOL OF OFFSPRING RESULTING FROM MATING NON-MEDULLATED AND MEDITJM-WEDULLATED EWES TO RAMS WITH NO,MEDIUM, AND HEAVILY-MEDULLATED FLEECES
Rain Ram, Medium- Ram, Heavily Means of Ewes Non-medu11ated Medullated Medullated Ewes
5* ß* R.P. S B R.?. S B R.P. S B H.P.
Non-me - dullated 0,00 5.00 2.50 0,70 21.70 11.20 9.60 76.00 42.8 3.43 34.20 18.80 Medium Medullation 0.40 14.60 7.50 5.50 58.30 31.90 48.50 96.50 72.5 18.10 56.50 37.30
Means of Rams 0.20 9.80 5.00 3.10 40.00 21.50 29.05 86.25 57.65 10.76 45.35 28.05
- Shoulder
B - Britch
R.P. -Reard1ess of position
'-J -J TBLE II. MEDULLATION IN THE OOL OF OFFSPRINGS RESTTLTIW) FROM MATIN LINCOLN x RÂMBOTJILLET ND ROMNEY x (LINCOLN x RÂMBOUILLET) EWES TO RAMS WITH NO, MEDIUM, AND HEAVILY MEDULLATED FLEECES
Ham Ram, Medium Ham, Heavily Means of Ewes Non-rriedullated Medullated Medullated Ewes
.P. s* B4 S B R.P. S B R.P. S B R.P.
Lincoln x Rarnbouillet O 12.07 12.On, 1.00 2ö.46l3.23 21.77 86.15 53.96 7.59 41.22 24.40
Romney x (lin coin X Rainboulliet O 33.24 33.24 4.1541.30 22.72 35.57 91.9? ]24; 55.49 34.36
Means of Rams O 22.65j2?.ö.5 2.5733.38 17.97 28.6789.03 58.85 iO.4148.35 29.38
*s _ Shoulder
B - Britch
R.P. - Regardless of position
I-J ewes produced lambs averaging 76 per cent medullation,
(Figure 7). Heavily medullated rams mated to ewes with medium medullation produced lambs averaging 96.5 per cent medullatlon. (Figure 8).
In tables I and II the medullatlon means of these lambs produced from the above mentioned different matings are presented. If we compare the pooled means, that is, the mean regardless of shoulder or britch, we find that non-medullated. rams produced offsprings with 5 per cent medullation, the medium medullated ram produced offsprings with 21.5 per cent medullat ion and the heavily medulla-
produced offsprings with 57.6 per cent medulla-
tion respectively, regardless degree of niedullation of
the ewes.
In the ewes of two different olassific8tlons of
medullation, the non-medullated. ewes produced lambs with
a medullation of 18.8 per cent while the medium medulla-
ted. ewes produced lambs with 37.3 per cent of medulla-
tion.
The lambs from the Lincoln x Ranibouillet ewes
have less medullation (24.4 per cent) than those from
Romney x (Lincoln x Ramboulliet) ewes (34.3 per cent).
1. Statistical Analysis
In order to analyze the results obtained by 14 mating rams of three classes or medullation with ewes with medium and no medullation, analysis of variance was used. The square-root transformation is applied to the original observed number of med.ullated. fibers for the pur- pose of using the analysis or variance which requires the measurement to follow the normal distribution (9, p.14). For example, if 16 medullated fibers are observed among 100 fibers, the measurement of inedullation is 4. This transformed measurement follows the normal distribution more closely than the original observations (9, p.41). The result of the analysis ol' variance is given in Tables III and IV. It is known that non-normality tends to announce too many significant results (9, p.24). In these analyses one percent significance level is used. for every test to counteract this tendency. The variance (Table III) due to differences in medullation of shoulder and britch wool indicates that there is a real difference between positions on the sheep. The offspring produced by the different mating combinations differed significantly. There was more inedullation in the fleeces of lambs from parents possess- ing raedullation than in the lambs from parents free from niedullation. Thus the variance associated with amount of medullation in the rams used shows statistical signif i- canoe. Also, the variance associated with the two 15
TABLE III. ANALYSIS OF VARIANCES OF RAMS OF THREE DIFFERENT DEGREES OF MEDULLATION MATED WITH EWES OF TWO DEGREES OF MEDULLATION
Variation Sum of Degrees Mean F Remarks Due to: Square of Square Freedom
Main effect:
Maie 586.82 2 293.41 84.60 *
Female 112.86 1 112.86 32.54 *
Position 496.37 1 496.37 143.11 * ist order interaction: Male x Fe- male 11.90 2 5.95 1.72
Male x Position 31.91 2 15.95 4.60
Female x Fosition 1.57 1 1.57 0.45
2nd. order intraction: Male x Female X Position 28.57 2 14.28 4.12
Error 374.58 108 3.46
Total 1,644.62 119
- Significant at i per cent 16
TABLE IV. ANALYSIS OF VARIANCES OF RAMS OF THREE DIFFERENT DEOEREES OF MEDULLATION MATED WITH LINCOLN-RAMBOUILLET EWES AD LINCOLN x RAMBOUILLET x ROMNEY EWES
Variation Sum of Degrees Mean F Remarks Due to: Square of Square Fr e e dorn
Main effect:
Male 725.90 2 362.95 65.55 *
Female 40.44 1 40.44 7.30 *
Position 703.97 1 703.97 127.13 * ist order interaction:
Male x Female 1.64 2 0.82 0.15
Male x Posi- 16.01 2 8.00 1.45 t ion
Female x 5.92 1 5.92 1.07 Position
2nd order interaction:
Male x Female 16.34 2 8.17 1.48 x Position
Error 797.35 144
Total 2,307.60 155
* Significant at i per cent 1? kinds of ewes shows statistical significance (Table III).
The first and second order or interactions were lacking in statistical significance (Thble III).
In Table IV is shown the analysis of variance for ewes of different breeds and for ranis of three dif- ferent medullation classes.
The Lincoln x Rambouillet ewes differed signi- ficantly from the Roinney x (Lincoln x Rambouillet) ewes in the amount of inedullation in the fleeces of their off- spring (Table IV). Again, the rams of three niedullation classes differed significantly in the amount of medulla- tion in the fleeces of their offspring. The first and second order interactions lacked statistical significance.
(Table Iv).
IV. DISCUSSION
It has been shown in the analysis of variance of the wool samples that differences between the three types of rains, two types of ewes, two different breeds of ewes and two different sampling positions are statistically significant at the one per cent level in both experiments.
It is believed that the genetical cause is likely re- sponsible for the medullatlon in wool, at least, for a large part. II1
From the table of means a clear picture can be
drawn that the occurrence of medullation in those lambs
are followed by the different degree of medullation of
their sires and dams. The heavily medullated. ranas mated
with medullated. ewes gave lambs iich are mostly nìed.ul-
lated. while the medium medullated rana and. ewes gave
lambs that are mostly medium medullated.
From the original data (counts on the medullation
per loo fibers, Appendix Table I) we found that the more
medullation the parents have, the more the lambs on the
average. At the same time, one or two heavily medullated
parents fewer fibers, while
most lambs from the non-medullated parents have certain
medullated fibers instead of none.
Parents with medium medullation have produced
lambs with about the same degree of medullat ion as those
parents, but some offspring with many and. some with few medullated fibers.
The results are of the kind that one would. ex-
pect in the light of breeding expression in general, and
it is evident that neither the non-medullated. condition
shown by the mating of non-medullated. rams to non-niedul-
lated. ewes nor the med.ullated conditions shown by the mating of heavily med.ullated. rams with medium medullated. 19 ewes behaves as simple Mendellan dominant. The data ob- tained from these matings, in fact, clearly indicate in- termed.late inheritance governed by multiple factors.
The absence of clear-cut segregation of offspring into definite and readily recognizable classes is typical of the inheritance of quantitative characters. In most of the studies of quantitative characters due to multiple factors, in animals and birds especially, no attempt has been made to determine the effect produced by single genes. It is, therefore, quite arbitrary to divide the rains and ewes into certain classes as medium inedullated and heavily raedullated, etc. That is why no attempt has been made to classify the lambs into definite groups according to degree of meduliation.
There is a very clear tendency in the results of this experiment for more medullation to exist in lambs than in their parents. This same situation has been observed in the New Zealand Romney sheep (18, p.157)
(8, p.T367). The explanation of this problem is quite obscure. The New Zealand scientists have been trying to find out the possible genes for the inheritance oÍ' hairy fibers in the Romney sheep for more than ten years. They postulated that there are three pairs of genes involved in the Inheritance of hairiness: one pair of dominant 20 hariness gaye one pair of recessive hairiness genes and. one pair 01' non-hairiness senes (4, p.552). This hypo- thesis agrees closely with the results of the cross be- tween rather low producing Holsteins and Galloways, which F2 animals have been obtained that produced. approximately twice as much. milk as the best P1 Holsteins (14, p.40). One of several explanations is that each of the P animals, besides carrying dominant genes for high production, also carried one or more recessive factors, differing in the two parents, for this character. As a result of recoin- bination in the :?2 generation, some animals would be pro- duced that carried recessive genes for high production, obtained from both P1 animals. One thing Is worthwhile to mention in the results is that two non-medullated parents produced a n:ethillated lamb. As to this fact one may recall the sanie occurence in poultry in the mating of white Wyandottes and white silkies in which pigmented offspring are obtained. This shows one of two things: (1) That non-niedullated. is doniinaiit and that me- dullated offspring cropping out is recessive In nature, or (2) That there are complementary genes involved such that both non-medullated parents are 21
contributing to a combination which Is
necessary for the expression of medullation.
There may be another explanation for the tendency
of higher percentage of medullatlon in the lambs than was
presented in their parents; the different sampling age of
ewes and lambs. The wool samples of the offspring were
taken as lambs, the age in which medullation Is higher
than in the second or third fleece.
In the study of the inheritance of medullation
between the two different oro ssbreed.s it has been found
that Lincoln x Rambouillet ewes produced lambs of less niedullatjon than those from Romney i (Lincoln x Ramboui1le
ewes. The reason is probably because the Romney breed has
a less evolutionary fleece, that is, more hairy fibers
in their fleece.
This investigation might be criticized because the same rani and ewe should have been mated for at least
two consecutive years ïn order to get two or more lambs
from. the same parents. And the F2 generation should be produced so that a much more aocurate picture of the in- heritance could be seen. Besides, the number of ewes used in this study were not large enough. According to
Miller's suggestion (16, p.119) 20 ewes should be pro- vided for each character to be studied. It is admitted 22 that this would have been the ideal procedure, but time and other conditions have prohibited a more extended study and besides, this study is only a part of the thole pro- gram in which a good crossbreed for mutton is being em- phasized.
Because there is an increase in the number of
Romney sheep in the Willamette Valley in recent years, it is worthwhile to discuss methods of examining and elimi- nating medullated fibers from the fleece. It is very interesting to know that in some of the high placing show rams they have quite heavy medullation in their fleeces4.
Therefore, it is advisable for the breeders to watch for this defect in their flocks and to select against this undesirable charact er.
The presence of med.ullated fibers in any wol is detrimental to quality from the standpoint of the manu- facturer. Barker (8, P.141) stated: "iuedullation is considered to indicate a hair and consequantly is anathema to the British manufacturer".
The medullated fibers are defective because of their hair character -- straight, coarse, and lacking of tensile strength. The spinning properties are lower, and
Nelson, O. M. Private Coxniiiunication. 23
In piece-dyed fabrics they produce a heathery effect by dyeing a lighter shade (2, p.132-133). The niedullated fibers are partially responsible for wide deviations from the mean diameter of fiber and unevenness in the fleece in which it occurs. Therefore the whole fleece will be graded lower. The only one tho thinks that medullation in Roniney sheep is of any value is À. 1. Barker, (5, p.646). He suggests that a new strain of Hairy Roniney sheep will produce the type of wool which is good for carpet manufacturing. Of course this is not the goal for the Roniney sheep breeders in the Wiilarnette Valley. While it is conceivable that factors extraneous to heredity are partially responsible for the formation of the medulla, the evidence thus far points strongly to the field of genetics. If then it is a genetic problem, the elimination of the niedullated fibers lies wholly in the hands of the breeder. If the problem. is to be solved the breeder should adopt a test for the presence of me- dullated fiber in the fleeces and of stud. animals and breeding ewes, especially in the yearling fleeces. It has been shown in the results that there is a difference In degree of inedullaticn between the po- sitions on the same animal. Results of this experiment 24 and those of coot's in New Zealand (10, p.46) indicated that britch sample will be good enough in representing the meduation of the whole fleece, There may be differences in the pattern factor for niedullation because some sheep show this only in the britch. However, a sheep with no britch nEdullation will show no medullation on the other parts of the body. Thus, one can use the britch sample to determine his animals that are free from niedullation and check the other parts of the fleece in. those animals that show britch medul- lation. While it is impossible for the breeder to do the sample testing, it is advisable to cull the ewes by checking their lambs at the age 0±' weaning. According to ble's (17, p.45) observation, there is a rapid decrease In percentage of niedullation of hairy-fleeced. lambs of the Rambouillet, Targhee, Corriedale and Columbia breeds, between two and five months of age. If at that time many medullated fibers exist, it is believed that these fibers will be of the persistant trpe and it will result in a meduliated fleece. The breeder could observe his lambs at this time and cull accordingly. Another test for medullation which is quite 25 simple and practical is the benzol test (20, p.T.68). It has been used in India and. New Zealand to a certain extent. It consists of examining a staple of wool by the naked eye while immersed in benzol, whichhasa refractive index practically the same as that of non-raedullated wool. The air-content in a medullated fiber, hast a refraction index higher than that of solid wool-fiber, enabling these medul- lated. fibers to be readily seen in the benzol by the naked eye, whereas, the non-niedullated. fibers are invisi- ble. Though it is not as accurate as Hardy's device, it Is indicative of niedullation and could be used by breeders special equipment necessary for the more accurate method. In addition, little technical training is needed to use the benzol method.
V. SUIv1MBY AND CONCLUSION
1. Rams with different degrees of medullation pro- duced. lambs with different degrees of inedullation accord- ingly. The heavily medullated. rains gave offspring with comparatively higher degrees of medullation regardless of ewes, etc. 2. Ewes which are free from medullatlon produced lambs with less medullation than those from iuedullated ewes regardless of rams. Since the percentage of xnedullation of all offspring are of intermediate type, therefore, a multifactor inheritance has been postulated. 3. Ewes of Lincoln x Rambouillet breeding produced lambs with less medullation than the ewes of Ronmey X (Lincoln x Ranibouillet) ewes. 4. Britch sample shows higher percentage of medul- lation than the shoulder sample either on the same indi- vidual or in all the samples. Therefore, the britch sample may be used as an index of the degree of ndulla- tion of the animal. 5. Elimination of raedullation from the fleece should be secured by testing the fleeces or stud ranis and breeding ewes and selecting accordingly. 27
1. Bryant, D. M. The incidence of kemp in the fleece of Scotish Mountain .blaokfaoe sheep with special reference to inheritance. Empire Tournal of Experimental Agriculture 4:165-185 April, 1936. 2. ergen, Vlerner von. American Wool 1andbook. 2nd enlarged edition, 1948. Textile 1300k Publi- shers Inc. N.Y. 1005p. 3. Dry, 1. W. The fiber types of New z,ealand Romney sheep. New Zealand Journal of Agriculture 48:269-277, Nov. 194.
4. Dry, E. W. A possible gene duplication in New Zealand Romney sheep. Nature 154:552. 1944.
5. Dry, F. W. Mendelian Inheritance in New Zealand Roniney. Nature (London) 160:646, 1947.
6. Dry, '. W. Hairy fibers of the Roniney sheep; IV. Fiber type arrays and hairiness. New Zealand Journal of Agriculture. 48:331-343, June, 1934. 7. Duerden, J. . Kemp fibers in the Merino. Journal Textile Institution. 17:T268-273, 1926. 8. Elphiok, B. L. The Detection and estimation of medullated fiber In New ¿ealand Romney fleece. Journal Textile Institution. 23:T36'?-T385, 1932. 9. Esenhat, Chmehlll. The assumption underlying the Analysis of Variance. Biometrics, Ìrch, 1944. 67p. 10. Goot, E. Hairiness in wook, Part I. Distribution of hairiness in the fleeces of New Zealand Rornney March ewe hoggets. New Zealand Journal Science and Tech. 27:45-Sd, 1945. 28
11. Goot, H. hairiness in vol, Part II. Yearly changes in hairiness in New Zealand Romney Iv.rch sheep. 27:173-187, 1045.
12. Galpin, Nancy. Tha occurrence of a britch-pool fiber-type array gradient In the New Zealand Romney lamb. Empire Journal of Exp. Agri. IV: 116-128, AprIl, 1936.
13. Hardy, J. I. Determination of fiber fineness. A rapid niethod using a new cross-sectioning device. Textile Research 3:381-387, 1933.
14. Ibsen, Heman L. Principles of GenetIcs. 2nd Edition, 1947, 118p.
15. thew, J. Merritt. The Textile Fibers. 2nd Edition. John Willey and Sons, N.Y. 1907. 48Op.
16. MIller, W. C. A general review of the inheritance of wool characters In sheep. Empire Journal Agriculture. 1:173-192.
17. Pohie, E. N. Keller, H. R., and Hazel, L. N. Mnthly changes In fineness, variability and medullation in hairy lambs. Journal of Animal Science 4:37-46, 1945.
18. Parr, Sir James. New Zealand sheep and wool. Wool Research and Textile World 35:157-160, 1929.
19. Rudall, K. M. Effect of shearing on hairiness in the fleece of the Romney lamb. New Zealand Journal of Agriculture 47:20-28, July 1933.
20. Wilson, J. :. The ndul1ated wool fiber. Hulgardia 4:135-152, June, 1929. .». s ¿
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.&PPFNDIX TABLE I. TI1J NUIYL3ER OF VEDULLAT.ED FIBERS PER loo FIBERS OF EACH OFFSPRING FROM THE SPECIFIED PARENTS
t, t t t E7ES " RAU t t, t
t t, ,, t t, t t t 't Free frein " Medium " Revy t t Ewe ' Medullation " Medullation " Medullation t t t, I, It t
t t Nuni-'t t ti t tt ? t e t S B* t? t ber " S ' B " S ' B t t t lt t I It t t
t t t, t tt t t t
t t 1 " o ' o o ' " o ' 95 t t, t t t, t ti t t
t lt t t t, t te t t
t t " 'e 2 o lo t 25 " 5 t 95 t t t t, t t, t t, t t
t o ,, H t tt t t, t t t -1 t 8 3 0 8 O ' 10 1 100 t t t, t t, t st t t
t it t it t t? t t t t 4tt ' o 3 O '60 " l 40 t t t t if t t, t i
t t? t it ti t t
t 0,5,1 0 t 5 ,, 4,85 60,l00 t t t It t if t t? t
0,10 0:20 2; 80 i t t 4 t ,, t is t 't _t
e t ti t ti t it t I t 7 t, O 12 t, O , 5 5 80 i t tt t t, t t? t t t t it t it t ft t t t t 8tt O i t, O 1 15, 50 t t t it t, t t, t t t ti t t, t t, i t t ,, , i O 2 5 60 g O t, t, , , t t t, t it it t t t i t? t t, t t, t i t tiO t' O O t, O, 4 t, 2, 60 t t it t it t t, t t S-shoulder, B-brltch kPPEI'1D IX T.BLE II. THE Ì'ItJMBER OF 1DULIAThD FIBERS PER 100 FIBERS OF ELCH OFFSPRING FROM THE SPECIFIED PARENTS
t t, - I " t EWES HAlvE I It t e
I ? yy T-r e ' e n Free rrorn Medium I, Heavy e ' It " t e Ewe Med.ullation Medullation Med.ullation e t-t ft e? t t e Nuxa-'t t t I 't t t ; 'ber" S' B " S B 5 ß , e et t It e te e
It t , : e tt t 15 20 ' 40 " 90 ' 100 t e It t t, t n e e t t t, t t? t t. T
" ' " 2 ' o .o 30 60 loo : ' t et t lt t t t, e t t re t t, t lt e t ' t 3" l'lO " 0'95 5 '100 e e It t ft t lt t t It t lt t lt e t e 't t 4 O ' 25 " 5 100 " 5 ' 100 e t n t t? 9 It t t t t, t lt t t? t e 9 9e " t lt 5 0 ' 7 O 60 100 ' 100 t t t t? t I? t n e
t lt t 99 t lt t '6" 0' 8 t? 0' 8 "lO'lOOe It t t lt t, t t
e te t ti t e 7 't 0 ' 15 " 10 ' 100 " 5 65 t lt t tI t t n t e
e ft t II t - '8" O'25 " 0' lO" ioo' loot t t t, e n t ev t t t t lt t lt t lt t t
t 9 o 6 " 5 t 100 15 t oo t e ti t it t lt t t t t t, t I? t I, t e t t tI t 3 ' 25 " O 40 " 95 ' 100 t t It t ti e n e t S-shoulder, B-britch 31
APPENDIX Tà.BLE III. THE SQ.UABJ R OOT OF TUE IVDULLÂTED FIBERS PER loo FIBERS OF EACH OFFSPRING FROM THE SPECIFIED PARENTS
I t? t EWES tt RAL t t n t
t t t, t, ft i i t n Free from " Medium " Heavy t , t, t ,Ewe Med.ullatlon t, Medullation " Medullation t It t, ti t t t t t, t ti t i, t Nu.ni t t ' t ber S B* S B " S ' B t t ,, t t, i i, t t t t I T-t T i T-T t
" i 2.23 0.00 ' 9.75 : " : ti , t? j t, t i e t T ti y t t TT- t t t ' 2 0.00 ' 3.16 " 1.73 ' 5.00 2.24 975 t T- et z , t-s t t t o t T H T " t T-T t t E-i' o.00 2.82 0.00 ' 3.16 2.24 ' 10.00 t t ;1 :: : t ¡.-4 t y .1 T!- t T
' z 1.7 0.00 7.75 " 1.00 ' 6.32 : 4 o.00 : ti e, , t :_. t t
t t u TI IT V t o ' " 2.00 ' 9.21 775 t z ei , 2.23 t t 1x4' te t, t t, t ?______T TT T-T t t t t, ' ,. t, t 0.00 3.lo 0,00 4q47 ' 1.41 ' 8.94 t 6 t t t, t e, t t t t TT ti y 'T 7 ? o.00 3.46 0.00 2.23 2.24 8.94 : : :: : :
t t -M- y TT t , ti t 7.07 8 0.00 ' 1.uO 0.00 ' 1.00 3.87 t t t t, t et , t, e
t T t It y T! VT t 2.24 775 ' 9 " 0.00 ' 1.00" 0.00 1.41 t t et ei te t t
t T t It y TI t tT
t t t " t " 10 " 0.00 0.00 0.00 2.00 1.41 7.75 t et t t t t et t tt S-shoulder, B-britch 32
A.PPE1DIX TABLE IV. THE SQUARE ROOT OF THE IVDULIATED FIBERS PER loo FIBERS OF EACH OFFSPRING 'ROM THE SPECIFIED PARENTS
, t EWES ' RALO ,, t I I VT T? t t t I, Free from " Medium n Heavy t t, t Medullation " Medullatlon " Medullatlon ,Ewe t ,, t, t t,
t t, t, NU- t t : ber S B s " S ' B t ,, :: t : : t t ti t t y t Ti.
' t 10.00 . 6,32 " 9.49 o.00 t : :: :: : : t t, t
t t I, r TT t , ,
t ' t ' 0.00 ' 3.16 " 2.24 t 5.48 775 , 10.00 2 t t It It t, t t t t t t t 1P
t? :: t t : 3.16 " 0.00 : 2.24 : 3 t t t, t, t t t t, t t t n 'I , , t t 't E ' 0.00 t 5.00 " 2.24 '10.00 " 2.24 t 10.00 t t ti t, t, t t t t t - t
t ;; t 0.00 : 2.65 " : 5 t I t 1t t, t e t
T t t ,, TI It t , t t " ' 0.00 ' 2.83 ' 0.00 ' 2.83 " 3.16 10.00 t t t t, It t e t t t t t, 9 t 9 t t t t 7 " 0.00 3.87 " 3.16 '10.00 " 2.24 t 8.01 I t t t. t? t e n t
t t t te t, t, t t t
t t, , , 10.00 8 0.00 , 5.00 0.00 3.16 l0.O0 t t t Tt ti t, t t
t t t t, t, t? t t
t et ,, 3.87 10.00 t 9 0.00 2.45 n 3.87 ,10.00 t t t t, t, et t t
t I, It te
, ,, 9.75 10.00 t 10 et 1.73 , 5.00 t 0.00 6.32
t t t t t, t, t, t
' S-shoulder, B-britch 33
APPENDIX TLBLE Y. THE NtTh[BER OF DDULL&TED FIBERS PER loo FIBERS OF EACH OFFSPRING FROM THE SPECIFIED PARENTS
t t? T
? EV1ES t RÀ.L t
, t tnt t t tt It t, t
i TT t Free from ' Ívledium " Heavy t t Ewe " Medullation " Med.ullatlon " Med.ullation t t ?t ,t 9 T t t Nuin_tt t ti t TI t T t " ?bertt s B " S B S ' B t t t, t t, t T, i t t t, t T, t t? t T t t itt o i tl t 95 It 5 t 100 I. S tt s. t t t t 17 tlOO ?t t t E-t 2 Q t i 5 5 95 t t. s e t t u. . - t t 3 ii o t o " o ' 20 't 98 ' 100 H e 1 t-. s e, t :D o , ' 1? t t , o t 15 t' O 5 15 100 . , . et . t t e. t çyP 5?t Q t 3 t' o t i tt95 t 100 y, t t e, te
t' n T ' 0 '80 O 5 5 95 ' ' t 6" T t
y, t t lo 3 30 15 50 o : :: : ,, i I t (.) t ti T? t ?t t t 8 8 0 0 1 90 , : ,,
t t T, lt t t 0 l2 o ,io 60 ,, :: t I
10 25 o 10 lo 90 :: : It : :: t T
t il 2 o 30 80 :: lt t t
t t ti i It t It T T 12 0 O 25 25 , ,
t t it t t, t it t t 13 ,, O O O , O O 65 ,
S-shoulder B-britoh 34
APPEI'IDIX T.kBL VI. TIlE I'flJMBER OF LDULIATED FIBERS PER 100 FIBERS OF EACH OFFSPRING FROM TIlE SPECIFIED P.BEW2S
t ? t? t EVT.ES 's RAJv t Vt
ti t, t t t t?
t' ti i t t n Free rrom t Medullation 't Medullation t t Ewe « Med.ullatlon , t, t, t t it
t? t t, t t t t Nnì_tt t
' t t t i: ' t S B t ber " S S B it e t? t T t t t, t 60 5 . : :: : : : - :: :: t o t it 0 60 1 40 í4 2 0 15 : :: t t- ti : ::
ti t t, i i t t k o o 60 , 100 ti ,, t t te t
t it t i t e ti t it
i t 95 t o , 8 ,100 100 t t i- t it t t t i-1 t it t t, '-4 i 100 5 o 70 4 t 85 , t t D t t, , ,
o t it t i t t Il t It 6 , 10 , 100 4 1 ,, O 80
It t ti t i i t ti t
ti O e 10 tto , 4 5 i t t
it t if t i s t it t
t t stO io s
t i ti t it t it t ,- t
- 5 ,, 15 95 , 90 1 o - O t o t i 40 80 90 H o 25 0 i-.1 I, t 60 100 t o 20 40 t t , : o 12 7 :o :90 t
T t, it it t t t_ t , 80 O 5 , i , 13 ,, O 100
+ S-shoulder B-britch 35
APPKND IX TABLE VII. THE SQ.UABE ROOT OF THE MEDULTATED FIBERS PER 100 FIBERS OF EÁ.OH OFJ?SPRIN FROM THE SPECIFIED PÂREIJTS
t EVES 't RA1v t ti e
It t t t, t? I t I et Free froni " Medium " Heavy t t Ewe " Medullation. " Medullation " Medullation t t It t, 't t t t Nu1n-" t It t t? t I t ft It " ber S J3 s ' B S ' B t t I? t t-t t ti t t t t 9 t 9 II t , , i ,, 0.00 1.00 , 0.00 9.75 ,, 2.24 10.00 t t It t It t lt t
, ,, 2 t-, 0,00 , 1.00 ,, 2.24 , 10.00 2.24 9.75 t i-1 t et t It t lt t t
, , t 3 ,, 0.00 , 0.00 ,, 0.00 4.47 9.90 10.00 t t t, t n I ti t I e t 4 it 0.00 , 3.87 0.00 2.24e 3.87 10.00 , :: t , 5 0.00 , 1.73 0.00 1.00 ,, 9.75 , 10.00 t t Vt t t? t it t t
,, , , ,, 2.24 , 9.75 t t 6 0.00 8.95 ,, 0.00 2.24
: , 0.00 3.16 ,: 1.73 : 5.48 , 3.87 7,07 t t.) t et t ft T n T t
8 t 0.00 t 2,83 ,t 0.00 t 0.00 n 1.00 t 9,49 t t i-1 t ti I II t
, , , t 9 tt 0.00 , 3.46 , 0.00 3,16 i a.24 7.75 t e te t lt t il t
, 10 , 0.00 , 5.00 , 0.00 , 3.16 , 3.16 9.49 t t ti t e, t te i t t 11 , 0.00 1.41 , 0.00 5.48 ,, 2.24 8.95 t t___ t, te t it i
, t t 12 t 0.00 t 0.00 , 2.24 , 5.00 t- 5.00 9.75 t t ii T it Ii t , 13 n 0.00 t 0.00,, 0.00 t 0,00 ti 0.00 t 8,06 e
T
S-shoulder B-b ri t oh 36
PNDIX TABI VIII. ThE SQUARE ROOT OF T1 EDULLATED FIBERS PER loo FI±3ERS OF EACH OWSPRING FROM TEE SPECIFIED PARTS
It t T
' , EVILS RAIVE t
, T? ,
1t ? ,t n t
ft t t ft Free from Medium Heavy 't t t Ewe ' Medullation Medullation ' Medullatlon , t T, lt TI t t t t t? t TI t j t t t t' ber S i3 S t B 't t B
t t, t t? t T t ;:; ti t t t!rl t ft t U tt t T
0.00 , 3.16 , 0.00 ".7 2.24 10.00 t t i 't ,
T t o , ,, t j t Vt t 2 0.00 3.87 OOO 7.75 1.00 , 6.32 t t ,, , 't t t ,t t TI t tt t t ,.4 , t t T, OOO , 0.00 , 0.00 0.00 , 7.75 10.00 t Ei t j, t ti t tt t T
4 , , t t t, o.00 , 9.75 0.00 2.83 10.00 10.00 t t t, t t? t ft t t 5 0.00 8.37 2.00 9.22 1.00 , 10.00 t o t , t m t t? t ti t it t t 6 ,, 0.00 8.95 3.16 , 10.00 2.00 , 10.00 t t , t t it t 't t it t t 7 0.00 3.16 ,, 0.00 2.00 2.24 10.00 t t ti t ti t It t
8 , 3.16 2.24 , 9.75 , 0.00 3.16 ,, 0.00
, 9 ,, 0.00 , 2.24 3.87 , 9.75 ,, 9.49 , 10.00
, t 10 n 0.00 t 5.00 t, 0.00 , 6.32 t 8.95 9.49 j t t ii t ti j, t t
, t t 11 t 0.00 t 2.24 4.47 , 6.32 7.75 10.00 t t t, t, t It t T
, 12 n 0.00 i 2.65 t 2,24 t 5.48sf 7.07 , 9.49 T t t Ti It t it t t
, 13 , 0.00 , 10.00 ,, 0.00 , 2.24 , 1.00 8.95
S -shoulder Bbr1 tok