ASEXUAL INHERITANCE IN THE VIOLET ( Viola odorata ) A RESIS PRESENTED TO THE FACULTY OF THE GRADUATE SCHOOL oF CoRNELL UNIVERSITY FOR THE DEGREE OF DOCTOR OF PHILOSOPHY BY ROY DAVID ANTHONY Reprinted from Technical Bulletin No. 76 New York Agricultural Experiment Station, March, 1920 # vs Rar : TECHNICAL BULLETIN No. 76. MARCH, 1920. New [ork Arcata J speriment Staton GENEVA, N. Y. ASEXUAL INHERITANCE IN THE VIOLET (Viola odorata). ROY D. ANTHONY. PUBLISHED BY THE STATION. BOARD OF CONTROL. Governor ALFRED FE. Smita, Albany. ComMISSIONER CHarues S. Witson, Albany. Irvine Rouse, Rochester. Frank M. Brapuey, Barkers. CHARLES C. Sackett, Canandaigua. Cuarues R. Meuuen, Geneva. Joun B. Mutrorp, Lodi. C. Frep Bosuart, Lowville. Parker Cornina, Albany. OFFICERS OF THE BOARD. CoMMISSIONER CHARLES S. WILSON, President. WituraM O’HANtoNn, Secretary and Treasurer. STATION STAFF. Wuirman H. Jorpan, Sc.D., LL.D., Director. tAuBerT R. Mann, A.M., Agricultural Economist. GrorGE W. CHURCHILL, Agriculturist and Superin- tendent of Labor. REGINALD C. Cotuison, M.S5., Agronomist. {T. Lyrriteron Lyon, Px.D., Chemist (Agronomy). James E. Menscuine, M.S., Associate Chemist (Agronomy). James D. Haran, B.S., Assistant Agronomist. Wituram P. WHEELER, First Assistant (Animal Industry). Rosert 8. Breep, Pu.D., {Witi1am A. Stockine, Jr., M.S.A., Bacteriologists. Haroup J. Conn, Pu.D., Associate Bacteriologist. Joun W. Bricut, M.S., GroraGe J. Hucker, M.A., Assistant Bacteriologists. Frep C. Stewart, M.S., tDonaLp Reppicxk, Pu.D., Water O. Giover, M.A., Associate Botanist. Mancet T. Munn, M.S., Assistant Botanist. Lucrus L. Van Stxyg, Pu.D., Chemist. Rupotpn J. ANDERSON, PH.D., {tLronarp A. Maynarp, Pu.D., Bio-Chemists. Botanists. ARTHUR W. CLARK, B.S., XIcHARD F, Keerer, A.B., Associate Chemists. Moraan P. Sweeney, A.M., Orro McCreary, B.S., Wiuu1am F, Wats, B.S., Assistant Chemists. Water L. Kup, M.S., Haroup L. Winston, B.S., Miuiarp G. Moors, B.8., Assistant Chemists. GrorceE A. SMITH, Dairy Expert. James D. Luckett, M.8.A., Editor and Librarian. Prerctvat J. Parrott, M.A., {GLenn W. Heretics, B.S.A., Entomologists. Huaeu Guaseow, Pu.D., *Frep Z. Harrzett, M.A. (Fredonia), Associate Entomologists. Rossiter D. Otmsteap, B.S., CLARENCE R. Purrrs, B.S., Assistant E’'ntomologists. Utysses P. Heprick, Sc.D., Horticulturist. tRouuns A. Emerson, Sc.D., Geneticist. {Wituiam H. CHanpuier, Pu.D., Pomologist. t Roy D. Antuony., M.S.A., *Frep E. Guapwin, B.S. (Fredonia), Orrin M. Taytor, GrorceE H. Hows, B.S.A., Associate Horticulturists, Wir C. Stong, M.S., Epwarp H. Francis, M.A., THEODORE E. Gary, B.S., Assistant Horticulturists. Jessie A. Sperry, Dzirector’s Secretary. Frank EH. Newron, Wiiarp F. Parcuin, Lena G. CurTIS, Mas M. ME tvin, Maups L. Hoean, K. Loraine Horton, Clerks and Stenographers. ELIZABETH JONES, Computer and Mailing Clerk. Address all correspondence, not to individual members of the staff, but to the New York AaricutturaL Experiment Station, GENEVA, N. Y. The Bulletins published by the Station will be sent free to any farmer applying for them. Connected with Grape Culture Investigations. pileeenbers of the faculty of the New York State College of Agriculture affiliated with this ation. { Resigned to accept position at Pennsylvania State College. i f : oe = sc TECHNICAL BULLETIN No. 76. ASEXUAL INHERITANCE IN THE VIOLET ( Viola odorata).! ROY D. ANTHONY. SUMMARY In the improvement of fruit varieties the question of f fixity of type in asexual propagation is of very considerable importance. The use of any of the tree fruits in a study of this problem would obviously extend the experiment far past the activity of a single investigator. In order to hasten work on this question the double violet, Marie Louise, which is propagated asexually, was used in a study of the effect of selection upon the length of blossom stem. Observations were also made of the inheritance of high and low yield. Four selection groups were made: long-stem plants of high yield, long-stem plants of low yield, short-stem plants of high yield, and short-stem plants of low yield. . The first year it was not realized how important a role plant vigor would play and so no record of this factor was made. Since then. the plants have been graded for vigor twice each season. One of the greatest difficulties encountered has been to find the best methods of showing the year’s work and of making the selections for the following year. The method of selecting the plants for propagation was changed after two years. Two sets of charts were used, one where the yield and stem- length of individual plants or of clonal groups are correlated with vigor and a second series showing the influence of location in the house upon yield, stem-length, and vigor. The length of all blossom stems is reported in one-half inch units and this unit is employed in the charts. Correlation tables are given for the three factors for the entire house for each year of the experiment and for each of the four selection groups for the last year. For purposes of comparison, the four peleciion groups were reduced to the same vigor by means of the regression coefficient. Environmental factors caused considerable variation within the same greenhouse, especially the first year of the experiment. The second year, the vigor and yield were approximately the same for the long-high and short-high groups but there was a lower stem-length average in the short-stem than in the long-stem selection. 1 Also presented to the faculty of the Graduate School of Cornell University as a thesisin partial fulfillment of the requirements for the degree of doctcrof philosophy. 3 4 This second year four plants were grown from every plant selected as a parent the first year. The record of these four plants showed that many of the first selections, based on the performance of one plant and without a knowledge of its vigor, were not correct. The third year the two high-yielding selections gave a slightly higher yield than the low selections but the difference was less than twice the probable error. At the same time in the two high- yielding selections the difference in stem-lengthin favor of the long- stem group was about five times the probable error and in the low- — yielding selections about nine times. | The fourth year there were supposed to be sixty-four plants from a common parent in 1914. In the long-stem selections, the high- yielding group averaged 3.108 + 0.441 blossoms more than the low-yielding plants. In the short-stem selection the high-yielding plants averaged 5.787 + 0.478 more than the low-yielding plants. In both the high-yielding and low-yielding selections the long-stem plants averaged, respectively, 0.361 + 0.036 units and 0.495 + 0.041 units longer than the short-stem plants. The fifth year of the experiment, in the long-stem selections the high-yielding plants averaged 1.847 + 0.754 blossoms more than the low-yielding plants. In the short-stem selections the high- yielding plants showed a gain of 4.970 + 0.767 blossoms. The long-stem groups in both the high and low yielding selections - gained 0.368 + .046 and 0.346 + .048 units, respectively. The process of selection has really been one of isolation whereby certain clonal lines have been selected out of the miscellaneous population purchased in 1914. We seemingly have proved only the existence of asexually inherited differences which probably were present before the experiment was begun. No attempt has been made to find when or how such differences arose. Tho the existence of such differences in the violet makes it seem more probable that there may be differences within a single variety of any fruit, the labor and the technical difficulties involved render it inadvisable for a nurseryman to attempt to find beneficial variations among fruits by bud selection. ' INTRODUCTION THE PROBLEM ~ All commercial fruits of America are propagated by asexual means. This fact creates a fundamental difference in the problem of the im- provement of our present fruit varieties as compared with the improve- ment of most vegetables and field crops where reproduction is by sexual means. g The development of the idea of pedigree and all it involves has had such a profound influence upon the live-stock industry that it a 5 is only natural for the fruit-grower to be influenced in his thought of his trees by this same idea. The growth of this trend of thought has been hastened by the exploitation of the term “pedigree” by a number of nurserymen who have sought by its use to convince the buyer that their trees were better than ordinary trees, tho the exact grounds upon which this statement was based seem frequently to be uncertain even in the minds of the nurserymen themselves. Since 1896, the Geneva Station has been working on a problem in orchard fertilization where the question of the fixity of type in asexual propagation is of considerable importance. That year an orchard of Ben Davis trees was planted for a fertilizer experiment. The trees were then top-worked to Rome, the buds all coming from a single tree. In 1912 a problem in selection within a clonal line was started when buds were taken’from the highest and from the lowest yielding trees in each fertilizer block and budded on own- rooted Spy stocks. These were planted on a uniform soil and are now nearly ready to begin fruiting. At about the time this second test was started, an attempt was made to study the fixity of type in the Baldwin by planting an orchard of trees secured from every part of the country and showing as wide a diversity as possible in their “pedigree.” It is too early yet to know what the value of this last experiment will be, but owing to the conditions surrounding the orchard it is doubtful if it can be relied upon to show conclusive results. It is obvious that work of this sort with species as slow in coming to maturity as are the tree fruits, must, of necessity, run far past the span of the working life of a single investigator. Even with the bush and small fruits progress would still be slow and these fruits are very susceptible to environmental changes. Therefore, in order to hasten the work on this problem, it was decided, in the spring of 1914, to grow the double violet, Marie Louise, in the greenhouse and to study the effect of selection upon the length of stem of the blossoms. That fall eight hundred plants were purchased from a commercial grower and planted in the greenhouse. The first question to be answered was whether from a mixed popu- lation types, or strains, could be isolated which would hold true to their selection year after year. The occurrence of “sports” in many horticultural crops is well known, altho when we consider the oppor- tunities for their production the number that have been isolated and proved to reproduce themselves is almost negligible. Some recent investigations would seem to indicate that citrus is an exception and that this genus is in a state of change, producing new types frequently. One variety of apple commonly grown in New York, the Twenty Ounce, has rather recently produced three sports, while a fourth can probably be credited to it. It is doubtful, however, if this number of sports has been found in all the other fruits grown in this State. Of course such sports represent sufficiently great changes to 6 ( be seen readily and to stand out from the minor fluctuations due to environment. Whether or not there are heritable differences too small to be detected in the commercial plantations is a question of fundamental importance to all engaged in the attempt to improve our fruits. As the first five years’ work with the violet seems to throw some light on this subject and as this year marks the beginning of the second phase of the problem, namely, whether by further selection _ isolated types may be shifted in either direction or even split into a multiplicity of types, it has seemed best to publish the data at this time. LITERATURE So many excellent discussions .of the general subject of bud varia- tion have appeared in the last few years that it would seem unneces- sary to present a very extensive bibliographic review. However, attention should be called to certain of these publications. The potato was the first asexually propagated plant in which improvement was generally sought by means of selection. A review of much of the early work with this plant is presented by Stuart (1915).” The results of one of the most successful experiments in the selec- tion of somatic variations were presented by Stout (1915). His work with Coleus did much to call the attention of investigators to this problem. Dorsey (1916) has given us an excellent review of the literature bearing on several phases of the question of bud variation. Jennings’ work (1916) with Difflugia is very interesting as it in- volves a quite different type of reproduction. The chapter devoted to bud selection in Babcock and Clausen’s book (1918) is a splendid critical study of the different phases of this problem. The most recent contributions from the horticultural standpoint have been Shamel’s articles (1918) dealing with citrus fruit improve- ment. DESCRIPTION OF MATERIAL USED The double violet, Marie Louise, for fifty years has been one of the most widely grown varieties in commercial and amateur houses. It is entirely sterile and is propagated by the separation of shoots from the original plant. So far as we have been able to find, the variety runs very uniform. While one or two so-called strains have been produced, the fact that they have never made their way into commercial culture would seem to indicate that they differed little if at all from the true variety. Altho the previous treatment of the 800 plants with which the experiment was started was not known, it ‘Is safe to assume that they would trace back to a very few parent 2 Reference to Literature Cited, page 30. u plants in a few generations as nurserymen are constantly propa- gating from their best plants and each healthy plant yields eight or ten cuttings. These plants were placed in the south house of a range of three running east from the main house. The south-east corner of the house is exposed to some of our coldest winds and fluctuations of © several degrees in temperature have been noted between this corner and the sheltered north-west corner. The main house at the west not only shelters from the wind but also has a tendency to decrease the light received in the north-west quarter of the house. This point will be discussed somewhat in detail later. Within the house are two tile-bottom benches, each holding 400 plants in fifty rows of eight plants to the row. These two benches will be referred to as the north and south benches, respectively. New plants are propagated each year either by allowing roots to form on the shoots, which are put out at the base of the crown, before the cuttings are taken, or the shoots are taken off just as the roots start to form. The shoots are then put in flats filled with sand. When root-growth is well started, the shoots are transferred to thumb pots and later, to larger pots and placed in cold frames for the summer. The plants are set in the benches about the first of September. The first few blooms that form usually have very short stems and are imperfect and these are discarded without measuring. The first regular picking is made about the middle of October. With a slight jerk the stems separate readily from the plant and the entire length from the base of the blossom is measured in units of one- half inch. As it was necessary to tabulate the results before plants could be selected for propagation for the following year, the last harvest record was usually made early in March, at a time when the plants were giving nearly maximum yields. METHODS House records.— Tno the experiment was to deal primarily with blossom-stem length, it was decided to study inheritance of high and low yield also as yield records had to be taken in getting stem- lengths. As plants were selected for long and short stem and high and low yield, this gave four selection groups: long-stem plants of high yield, long-stem plants of low yield, short-stem plants of high yield, and short-stem plants of low yield. In order to shorten the records and the discussion, these groups are usually referred to as follows: long-high, long-low, short-high, and short-low, respectively. The records are taken in the greenhouse on large sheets 14 by 17 inches, cross-ruled into quarter-inch squares. The plant number is placed at the left. and the date at the top. Each flower-stem is measured and the length to the nearest one-half inch placed in the 8 next square. When each harvest is over, a vertical line is drawn outside of the last record of the highest yielding plant. In this way the first records of the following har vest for all plants are in the same ’ vertical line of squares, making it easy to study the yield fluctuation of any particular plant from harvest to harvest, and showing the relation of any plant to the maximum yield of that harvest. With conditions at their best, the bed is picked over every week or ten days but in very’ cold weather it may be three weeks or more between harvests. When the large record sheets are filled they are totaled for each plant. In an average season, when the third series of sheets is filled, the records are discontinued ‘and propagation com- menced for the following year. For the first year the various factors were studied for each of these three harvest periods but since then only for the total harvest. When the experiment was begun it was not realized how important a role plant vigor would play and so the first year no record was made of this factor but the need of it was clearly seen when it became necessary to select plants for propagation for the following year. ‘There seemed to be no feasible way of weighing the plants or measur- ing their leaf surface and so the vigor was estimated in percentage, the most vigorous plants being graded as 100 and a few plants, with only a half dozen or so leaves and practically no blossoms, graded as 10. Two persons cooperated in this work, each checking the other, and it is doubtful if a third person would have shifted any of the plants more than ten points in the vigor score. The vigor was taken twice, once when the plants were coming into full bloom early in the winter, and a second time just before the selection of the plants for propagation for the next year. In computing final records the average of the two observations was used. In a very general way, plants recorded as having a wanna vigor were somewhat alike from year to year; but, because of the varia- tions of the plants in different seasons, it is not possible to compare _ the vigor records of two different years with each other except in so far as each is considered as a statement of the relative values of the plants for that particular year. Statistical methods.— In order to simplify the problem of studying the plants and especially of making selections for the next year’s propagation, the number, yield, and stem-length of each plant was placed on a single card and the cards grouped according to the line of selection. ‘These cards could then be shuffled for any factor, and frequency charts and correlation tables very quickly prepared. This method has resulted in a considerable saving in time and has-been a very important help in analyzing the data. One of the greatest difficulties encountered in the experiment was to find the best method of showing the results of the year’s work and of making the selections for the following year. The method of selecting the plants for propagation was not “the same for the first 9 two years as is now used. In these early years the records were thrown iato frequency tables and certain maximum or minimum yields and lengths selected until the number of plants which it was planned to propagate was isolated. This is well illustrated by Chart I which shows the selection standards for the first year in connection with the frequency curves of length and yield. The lines from which the arrows are drawn show the selection limits. Cuart I.— FREQUENCY Curves oF LENGTH AND YIELD, 1914-15. LENGTH. ; YIELD. 100 ect Leng fi IS y My g si eS 2) . J S fo) = S ©) ‘= a 2 ts) 2 a = ie.) for) S Length in one-half inches. Yield per plant. By selecting the plants from the frequency curve and without knowl- edge of their vigor, any plants which were high-yielding because of ab- normally high vigor were selected and the same was true with plants of low yield, due to lack of vigor. This objection has now been overcome by a somewhat different method of selection. Each plant or group of plants is plotted according to vigor on a chart with a common base line and with the yield plotted above and the stem- length below the vigor line. (See Chart IV.) 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As an aid in selection, a straight line was drawn thru each population in such a way as to divide the members in the various vigor classes into nearly equal parts. Inasmuch as the individuals near this line, the average plants, were discarded when- ever possible, this method of dividing the population was deemed sufficiently accurate. To check this point, however, in several cases the straight line was determined by the formula, y=mx-+n. The two lines were found to differ but little. Values lying above the line would then represent high-yielding or long-stem plants and_ those below the line low-yielding or short-stem plants. By selecting from this chart it was possible to find those plants which were above or below the average at any particular vigor. This method has been used the last two years with very satis- factory results. In 1915-16 the selections were made on the basis _ of the average performance of the four plants tracing from a common origin the previous year. The following year the average of the six- teen plants with common origin was used and so on for each year, the total number tracing from a single plant in 1914 increasing by mul- tiples of four each year. A second series of charts (Chart III) is used to show the influence of location in the house upon the three factors studied and the rela- tive fluctuation of those factors. These charts are also useful in showing the sudden jumps that frequently occur in passing from one type of selection to another when the plants lie in adjacent rows. Correlation tables are made for the three factors for the entire house and for each of the four selection groups, and for each of these the following values are computed: Coefficient of correlation of vigor and length, vigor and yield, and length and yield; the standard deviation of length, yield, and vigor; the mean values of length, yield, and vigor; regression coefficients; and the corrected means where yield and length have been reduced to a common vigor by the regression coefficients. These values are summarized in Table I and in the appendix are placed the correlation tables for the entire house for each year of the experiment and the correlations for each selection group for the last year. To include all the correlation tables seemed unneces- sary since those which were selected show the general trend of the correlations. A study of the correlation tables themselves without reference to computed values is of considerable help as it shows roughly the degree of correlation, the approximate averages, and the presence of any abnormal plants which need further study. If any sporting occurs in the factors studied it can be quickly detected by this means. In studying the 1915-16 records a number of the correlations were worked out for the first and for the third periods of harvest. 13 The correlation between vigor and yield in the first group of pick- ings was low but for the third group considerably higher except for the short-low group and in the total this correlation is still higher 2dydy NOySy’ shows the mathematical reason for this. The total yields for the third group were of course considerably higher than for the first group of pickings and the total for the three groups very much higher. Thus, we have a constantly increasing *dd in the numerator, while in the denominator the standard deviation for vigor does not fluctuate materially and the standard deviation for yield does not increase as rapidly as the summation so that we have a constantly increasing value for the coefficient of correlation. The fundamental reason is probably that by the third period of harvest the plants have reached their maximum production and under such conditions prob- ably show greater correlation than earlier in the season. This varia- tion in the correlation is another reason why it is difficult to compare different years with each other. ‘i In Table II are summarized for the last four years the corrected mean yield and mean stem-length for the house and the differences for the contrasted selection groups. Such a summary is very con- venient in the analysis of the data. with all groups. >-+---- Stem-length. ‘ o _——< oe Vigor. 70 8.25 21 60 8.00 18 50 7.75 15 40 7.50 12 30 7 25 9 7.00 6 . Rows 10 29 30 40 50 \ 80 8.50 24 60 8.00 18 SourH SIDE. 40 7.50 12 20 7.00 6 Rows 10 £0 30 40 50 Cuart VIII.— Rea Ave. yield per plant. 25 15 xg) Vigor 20 30 40 8.50 8.00 7.00 Ave. length =19: ined e blia e iae Py tad Eto Weyigetene hee 1 Cuart VIII.— RELAtion oF YIELD AND STEM-LENGTH TO Vicor, 1917-18. Record of 12 adjacent plants. Ave. yield per plant. ry é $ 25 $ ¥ ¥ 4 e w $ a 20 8 g B. P Ps * « £ y - wt = ™ Ld " = 15 = u a Pd Af g = g “go x* ¢ 9 g 10 “es x * g 4 S Selected for high yield. P Selected for low yield. 3 8.50 3 > x ry 3 r a $ 4 8.00 & - e x v a {. a ¥ um 1 a x = y 7.50 : + §¥ * « «A : g A g C) x x 2 z : « = 3 Selected for long stem. 7.00 Y 2 ¥ « g Selected for short stem. Ave. length :) wey eae) Aare Bac ih? bo Ou Crarr [X.— INFLUENCE oF LocaTION IN House ON YIELD, STEM-LENGTH, AND Vicor, 1918-19. 4 Averages of 8 plants. gl Bus KOE NoRTH BENCH. 50 a os Peal oO - Sak et 95 9.00 45 NORTH SIDE. Yield. ----- Stem-length. 90 8.50 42 ae ees VIZOF. \ 6L)- Retained for 1919-20. . 85 8.00 39 80 7.50 36] 75 7.00 33 70 6.50 30 65 6.00 27 be <— tee. Rows 10 20 30 40 74 EF 90 B 51D 95 9.00 90 8.50 SouTH SIDE. 85 8.00 80 7.50 Wy 00) 70 6.50 65 6.00 Rows : 10 20 30 40 74 50 E 29H > Vigor. 90 80 70 90 80 75 65 60 26 Cuart IX (Continued). SouTH BENCH. Norra SIDE. Yield. Ssooos Stem-length. 8.50 42 ee Viror. tained for 8.00 39 7.50 36 7.00 33 6.50 30 6.00 27 5.50 24 Rows 10 20 30 40 9.00 45 8.50 42 8.00 39] SoUTH SIDE. 7.50 36 7.00. 33 6.50 30 6.00 27 “Rows 10 20 30 40 90 B 51D OE Cuart X.— ReLation Ri Ave. yield. per plant. 51 48 45 39 36 33 30 27 24 Vigor | 63 70 15 8.90 8.60 3 8.00 7.70 yw 6.80 g Ave. length in 4 inches. Cuart X.— RELATION or YIELD AND STEM-LENGTH To Vicor, 1918-19, Records of 8 adjacent plants. Ave. yield- per plant. $ R 4 ah : Yield. & $ ¢ @ g g $ 3 Pe oY) 3 g 4 g $ % q g ; y 5 3 j 2 g SO High 3 g 9 e a ig oe ag e° 99. gre ge 29 ; $ § oo g g a On eee 80 85 90 95 $ $ a 8.90 3 4 $ vat k $33 33 8.60 boas enn 5 ly ; y ; 8 C3) $ 3 2 g me} 8.30 Seeatitutrous on ' 4 a 0 y Length. 3 4 4 $ g 8.00 g nA i yen 8 e ¢ ao) 7.70 2 3 ast $3 $ g 7.40 ¢ ae AS ee 7.10 6.80 g Ave. length in inches, ; Ta): ae ay Cube jue yee i aN " erie Wen RUNS ie Re aia ey Hah me att 27 short-stem blossoms but, on account of the large number of blooms in the first harvest this year, it was decided to retain it in the records. Because of including these short stems, the average length has prob- ably been decreased several tenths of a unit. In the third group of harvests, that extending from December 24 to January 31, a very considerable proportion of the plants averaged well above ten units in length. The average vigor this year was 81.447 + .339. In other years it has been the custom to arrange the vigor scale in such a way that the average would be approximately 50, but this year such a large proportion of the plants showed extreme vigor that this did not seem advisable. Plants recorded as having a vigor of 50 per cent this year were probably fairly comparable with those having a similar vigor record last year. Group yields.— In the long-stem selections, the high-yielding plants averaged 1.847 + .754 blooms more than the low-yielding plants. (See Table II.) This is less than the difference between the same groups in the previous year. This fact may probably be considered as another indication that our process of selection is beginning to isolate distinct groups because this year there are progeny of only two of the original plants represented in the long-high group. One of these plants, 51 D, which is the clon retained to continue the selec- tion for another year, is not strictly a high-yielding clon, but, from its performance of other years, would be graded as somewhat above the average. It is retained, however, because it is long-stemmed and is sufficiently high-yielding to answer the purpose. In the short- stem selection the high-yielding plants gave 4.970 + .767 blossoms more than the low-yielding plants. This great difference in yield is due to the fact that 90 B, the short-high strain, is a remarkably high-yielding group. Group lengths.— In the high-yielding strains the long-stem plants were .368 -- .046 units longer in blossom stem-length than the short- stem plants and, in the low-yielding strain, the difference was .346 + .048. (See Table II.) These differences are not quite as great as in the previous year but, as before explained, this is due to the inclusion in the records of a large number of short-stem blooms in the first picking which in previous years were discarded. When the house was planted in the fall of 1918 the different lines were broken into groups, the groups being placed in different parts _ of the house. In all there were twelve groups. A study of these in Chart IX shows that in seven cases the relative position of the graph of both stem-length and yield indicates clearly the type of the selection of that group. In each of the other five cases, the graph of either stem-length or yield is in the position to be expected from its types of selection while the other graph is either uncertain or apparently opposite to the selection. In two of these latter cases, the two groups of 74 F on the west end of the north bench, a possible 28 cause for the failure to follow the selection may be suggested. If we refer back to Chart II we see that with the mixed population of 1914-15, in the area occupied by these two groups, there was a dif- ferent relation between the stem-length and yield lines than in the rest of the house. As has been suggested before, this difference is probably due to a slight change in the environmental factors in the northwest corner of the house. The way in which the plants adhere to their respective types of selection is also shown in Chart X. Here the whole population is shown in groups of eight plants and the character of the selection of each group is graphically indicated. It is remarkable the small number of groups which fall in areas occupied by groups of the opposite selection. CONCLUSION The first five years of selection have isolated four well-defined groups. The differences between the opposite selections are suffi- ciently large to show graphically on the charts of the two benches for the fourth and fifth years and, in comparison with their probable errors, make it certain that they are not due to chance variations. It is believed that variations due to lack of uniformity of conditions within the house have been so nearly eliminated that the final results are not materially affected. When this work was started it was the opinion of those connected with it that such a result as this would not be secured and probably most pomologists would have held the same opinion. For this reason there is justification in restating certain points and showing the possible application of the results. During the five years reported in this publication, the process of selection has really been one of isolation whereby certain clonal lines have been selected out of the miscellaneous population purchased in 1914. In nearly every case each plant within the clon has been the parent of four plants used the following year. It follows, then, that we seemingly have proved only the existence of asexually inheri- ted differences which probably were present before the experiment was begun. No attempt has been made to find when or how such differences arise. That differences have been found in the violet which could be passed on from bud generation to bud generation does not prove that similar differences may be found in the apple but it does make it seem more probable that such differences exist. Unfortunately, from the standpoint of practical application, the labor and the tech- nical difficulties involved in proving that an observed difference is really transmissable and not simply a temporary response to an environmental change make it seem inadvisable for a nurseryman to attempt such a problem. 29 The four selection groups now contain only five pure lines, each tracing back to a single plant in 1914. Whether these clonal lines are pure lines in the sense that Johannsen has applied that term or whether they are simply stages beyond which we will go to greater differences, thru further selection, is now the problem. PLANS FOR CONTINUING THE EXPERIMENT There had been no selection within any clonal line up to the spring of 1919. At that time it was decided that fifty parent plants would be saved in each of the four types of selection as this would give the 800 plants necessary to fill the house. In all but one type these fifty were selected from one clon only. As there were from 100 to 150 plants from which to select the fifty parent plants this afforded an opportunity to choose those plants which best answered the conditions of their particular type. Instead of selecting single plants, groups of four, each tracing back to a single parent the pre- vious year, were selected. In one group, the short-high selection, two plants were used to start two new lines in an attempt to break up a clon by selection within it. One of these plants was selected for long stem and the other for short stem. In choosing these, care was taken to find two plants which did not have a common origin until the original 1914 plant was reached. These two lines will be continued with intensive selection. These general plans will be followed for another five years to test the fixity of the clonal lines. ACKNOWLEDGEMENTS Thruout the five years of the experiment Mr. Joseph Welling- ton has had the oversight of the propagation and the harvesting. This has involved a great deal of work and no small part of whatever value the work has is due to the care and accuracy with which the records were taken in the greenhouse. He has also assisted in the preparation of the data for publication. The author is greatly indebted to Dr. H. H. Love of the Depart- ment of Plant Breeding of the New York State College of Agriculture for many helpful suggestions and advice in the biometrical work. The initiation of this work was due to Dr. U. P. Hedrick. For many years he had been interested in the question of pedigree in fruit and saw the possibilities in extending this study to some plant where results could be secured more rapidly than with the apple. His direction has been one of the factors in the success of the work. 30 LITERATURE CITED Stuart, Wm. 1915. Potato breeding and selection. U.S. D. A. Bul. 195. Stout, A. B. 1915. The establishment of varieties in Coleus by the selection of somatic variations. Carnegie Inst. Wash. Pub. 218. Dorsey, M. J. 1916. The inheritance and permanence of clonal varieties. Proc. Am. Soc. Hort.Sei. 1916. Jennings, H. 8. 1916. Heredity, variation and the results of selec- tion in the uniparental reproduction of D jfflugia corona. Genetics, 1 No. 5. Babcock, E. B. and Clausen, R. E. 1918. Genetics in relation to agriculture. McGraw-Hill. New York. Shamel, A. D., et al. 1918. Citrus fruit improvement. U.S. D. A. Buls. 623, 624, 697. | 31 “Our J[Vy-9u0 fo SYIUN UI WOAIS ov SYYSue] UIE UOssoTg | ‘uo1ye[ndod Sutpusose ue Aq UMOYS SI UONVIILI00 dATyIsod B sty} Jo asnvoaq puR 9[qe} 9Yyy Jo: doy ayy 4B SON[BA JSOYSIY ItoY} YIM polpNjs si10zoVj oY] OSULLIC 07 JUSTUSAUOD B1OUI UVEG SeY Ji SaTqe} UOIZV[III0N OY} UT y Gl) SAS Se! 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Total for entire house, 1918-19. STEM-LENGTH. 45 a =| x IDIDDHDaAOMr~-ONMONnAOOOMMH 5 ee OS 68 RIGS tO RO DISA i= : Fs cl Il OL G + 166 . mH a aH rH 126 : N ANMMON MRA AAS 1¢°6 o re COMIN N AH TNN mo TE 6 0 mo — MBSEOMIMDAOANIO WHMAN Ss IL '6 O me SH HH oO CO SHLQTHHN NAN SG 16°8 9 MAL OI DOWN O19 SF IZ’8 = 1 nN . AMMAN MMO OmMN MMR OMHAN MN T&°8 es * SANTOR RH ONOMHN Ils ee O MPSeH COND OmO FR OMOn SS 16°2 4 c MANS MMOMOOIMO OMAN 4 a, 4 . Mena We bh 192919 19 MS 1g 2 0 mal CN rt OD OD OD OD Pe 0 10 OO N T§ 2 5 Se Nts EZ 5 N mN ANN 169 . re N 1Z°9 . re T¢°9 . —_— Ig 9 Mm « aO S HHOLONH OO Or Heo 28 ROBSSS SO STAHAOHMONANA oS Om € 22 o9 83. 88° 681 71 «41 4643 Dab lOe 38.4956 1 TOTALS. r= .379+.021 44 2 080° S79" =: I. 1 OU 8 AG ASS AKG cS tells ts} 5 XG! 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