A STUDY I or SOME-TAM CHARACTERS“: 7 i ’ orfsmoom {-15sz BARLEYS ‘ ‘ THESISFUR THE DEGREE, or M. s; C. Roy Adair» 1932 A STUDY OF SOLE AWN CE‘LRACTERS OF SMOOTH- AWN BARLEYS ACKNO WI. EDGLLEJNTS The writer is very grateful to Professor E. E. Down for the help and guidance he has given throughout the course of this problem, and for his and Er. H. M. Brown's constructive criticism in the final review of this paper. A STUDY OF SOLE AWN CHARLLCTERS OF SHOOTH AWN BARLEYS Thesis Respectfully submitted in partial fulfillment for the degree of Master of Science at Michigan State College of Agriculture and fipplied Science 0. Roy Adair 1952 THE‘A‘S II III IV VI VII VIII IX TABLE OF CONTEHTS Introduction Reasons for studying this problem ........... Statement of problem ........................ Review of literature .......................... Source of material ............................ Mbthode ....................................... Factors studied ............................. Description Of terms ooooooooooooooooooooocoo 'OCDmCDQCfll-‘H Computations .................................. Results ....................................... 10 Discussion .................................... 21 Summary ....................................... 25 Literature cited .............................. 27 Appendix ’ Tables 1 - 17 Figure l Plates 1 - 4 Tables 18 - 25 93754 IHTRODUCTION comparisons made at the Michigan Agricultural Experiment Station in 1928 and reported in Michigan.Agri- cultural Experiment Station Special Bulletin No. 191 indicated that there were pronounced differences in the smoothness of awn of "smooth awn" varieties of barley {10). This condition raised a question as to the possibility of obtaining six-row or two-row strains with awns of still greater smoothness. In 1929 a problem was outlined with the idea of obtaining some information.on the inherited and seasonal variations in the degree of smoothness which might be found in the smooth awn barley strains then grown on the Station plate. The earlier work at the Station on smoothness of awn had shown that smoothness is not a simple quantitative character. With this in mind.five factors were studied: length of awn, length of barbed area, ratio of length of awn to length of barbed area, and the number of barbs in areas one centimeter long taken at two different places on the awn. The main objects of the problem were as follows: 1. To determine whether it is as reliable to use strain averages as to use individual head averages in the calculation of results. 2. To determine whether there are significant in- herited and seasonal differences between the degrees of smoothness of awn or the lengths of awn of the six-row strains and those of the two-row strains. 3. To determine whether there are significant in- herited and seasonal differences among the degrees of smoothness of awn or among the lengths of awn of the six- row'strains; of the two-row strains. 4. To determine whether there are significant in- herited and seasonal differences in the degrees of smooth- ness of awn or in the lengths of awn of a group of two-row strains, all of which come from the same cross. 5. To compare Michigan Black Barbless, Spartan, and MichiganeTwo-Row with some of the extremely smooth and the extremely rough strains. 6. To determine whether there are significant differences among the variabilities of the different factors studied. 7. To determine the degrees of relationship existing among some of the factors studied. The literature bearing upon the different phases of this problem will be reviewed before discussing this problem as outlined. REVIEW OF LITERATURE Harlan and Hayes (4): Crosses were made between Lion and rough awned varieties. They determined that a high yielding, smooth awned variety, of any head type de- sired, may be produced by crossing and selection. Harlan (5): The term "smooth awn" may be confused with awnless or hooded. Smooth awn barley was first described by Koeraricke, EurOpe, in 1882. Robert Regel, Russia, worked on smooth awn barley in 1909. The United States Department of Agriculture and the Minnesota Agri- cultural Experiment Station started a cooperative project on smooth awn‘barley in 1909. An importation of smooth awn barley was made from North Africa, in December 1911, and crosses were made between this smooth awn barley and common rough awn varieties. Smooth awn was found to be recessive to rough awn. This author had no conception of the number of factors concerned so could not say whether teeth can be entirely removed or not. He stated, however, that Hordeum lieorrhynchum (smooth awn types) importations have yielded as well as other importations, and smooth awn progeny have yielded very well in nurseries, but the absence of smooth awn commercial varieties in Asia and southern Eur0pe Where they have been known for a long time ;indicates a weakness of some sort. Indications are that smooth awn varieties that give satisfactory yields may be obtained. Harlan and Pope (6): The smooth.awned character of Dion is undoubtedly linked with undesirable characters, but this linkage is not absolute, as it has been found that crossing over will occur. When back-crossing is practiced the chromosome carrying the smooth awn factor will cross over with the chromosome carrying rough awn factor and desirable characters, so there would be a maximum chance of smooth awn becoming linked with desir- able characters. Hayes and Wilcox (7): The work done by these authors showed that the smooth awn barley overcomes the undesirable characters of the rough awn, awnless, and hooded varieties. The awn is an important physiological organ and the lack of it in the awnless and hooded varieties causes them.to be low in yield. The rough awn of the "rough awn" varieties makes them very disagreeable to handle. The smooth awn varieties are not disagreeable to handle and they do not have the physiolOgical limitations of the awnless and hooded varieties. Vavilon (12): This author worked on the origin of smooth awn barleys and he believed that they are the result of natural crosses between two rough awn barleys of different types. Types of rough awn parents necessary to get smooth awn progeny are: slender awn with triangular, wide-apart, acioular teeth arranged in close spiral (group nutans colchicum and n. precocius), and broad awn with large closely set teeth, disposed in loose spiral and occuring also along the external median line of the awn (coeleste and nudoficiens groups). He believed smooth awn is dependent upon five or six factors. Colin and Trouard-Riolle (l): Crossed white, rough- awn.Albert and smooth-awn black barley. Some of F1 heads had smooth awns only, others had smooth awns and rough awns and some others were smooth for half their length and rough for the remainder. Only the rough awned heads showed any signs of Mendelian segregation in F2. Her (9): This author worked on the interrelation of the following genetic factors in barley: black glume (B), rough awn (R), long rachilla hair (L). The arrange- ment is BRL, the distance between B and R being greater than the distance between.R and L. Crossover value between B and I. is 44.04“.“55, between B and R is 41.48i5.45% and between R and L is 34.512.89%. Griffee (3): Svanhals, a two-row, rough awn, White glume, Helminthogporium sativum resistant variety, was crossed with Lion, a six-row, smooth awn, black glume, H. sativum susceptible variety. The F1 was rough; F2 rough and partially or almost entirely smooth. The results gave indication of a two factor difference, RR and SS. R produces rough, S is hypostatic to R.and in absence of R produces intermediate type, rr 88 produces smooth awn of the Lion type. There is quite a variability in the placing of the barbs on awns of pure lines, which indicates the presence of other factors. In.the F2 he obtained twelve rough, , three intermediate, and one smooth. Rough and smooth awn factors are independent of two and six-row, and black and white glume. One of the three factors that produces resistance to H. sativum is linked with rough. Hayes (8): The first series of crosses made by Dr. H. V. Harlan between Lion and rough awned varieties failed in Minnesota because of susceptibility to Spot blotch (H. sativum). A second series of crosses was made and the problem of spot blotch resistance was taken into consideration. The factors for rough awn and re- sistance are linked but some crossing over occured so it was possible to obtain six-row, smooth awn, white glume, resistant varieties. Velvet (Minn. No. 447), Comfort (Minn. No. 451), and Glabron (Minn. No. 445) are the three best varieties obtained from this series. Sigfusson (ll): Crosses were made between Chinese, a rough awn short rachilla hair variety, and Lion, a smooth awn, long rachilla hair variety. Rough awn was found to be due to two complementary factors, R and S: R being more important than S, but both necessary to produce roughness of the Chinese type. There was some linkage between r and L, the factor for long rachilla hair. Rather, Down, Brown, Clark (10) P.18-l9: Photo- micrOgraphs showed differences to exist in the degree of smoothness among several of the commercial "smooth awn" varieties. Spartan and Michigan Black Barbless were the smoothest of the varieties under observation. SOURCE OF MATERIAL Three commercial varieties, Michigan Black Barbless, Spartan, and Michigan Two-Row, and some 222 strains coming from crosses made in 1924 between the smooth awn and the rough awn or hooded varieties in the nursery at that time are the material used in this thesis. The three varieties were used for comparison purposes and are some of the parents of the smooth awn strains. Michigan.Black Barbless is a six-row, smooth awn variety that was selected from Lion (C.I.No.923); Spartan is a two- row, smooth awn variety and is a selection from a cross be- tween Michigan Black Barbless and Michigan Two-Row: Michigan Two-Row is a two-row, rough awn variety. There were 76 six-row smooth awn strains coming from five crosses and 144 two-row smooth awn strains coming from twelve crosses. These 222 strains had been selected for four years until they were breeding true for apparent morphological characters. During this period of selection the smooth awn segregates had been saved and the rough awn segregates discarded. The smoothness had been judged by pulling the awn back and forth between the thumb and forefinger. HETHODS Ten heads were collected in the field from each of the 222 strains and the three varieties and placed in an envelOpe. Four awns were taken from each head and the following measurements and counts were made: length of awn recorded in millimeters; distance the barbs extend down from the tip of the awn, recorded in millimeters; number of barbs in an area one centimeter long starting .5 mm from tip of awn (called"first count° in this thesis); and number of barbs in an area one centimeter long start- ing 25 mm from tip of awn (called”second count”in this thesis). A fifth factor, the awn-barbed area ratio, was obtained by dividing the length of awn by the hength of the barbed area. Three different methods were used during the course of this problem in making the counts and measurements. Early in 1929 a bifocal microscOpe was used but it proved unsatisfactory as it consumed too much time. A change was then made to a stereOptican. The awn was placed on a graduated slide and the image projected on a screen with the stere0ptican. The awn was thus magnified so that it was possible to count the barbs. This method was found very impractical in.that it took two men to Operate the madhine and make the reading, so a projection microsc0pe was obtained to use on the 1930 material. The magnified image of the awn was projected down on the table, thus making it possible for one man to Operate the instrument and make the counts. COLIPUTATI ON S The results obtained from the two and six row strains wmre computed separately. Two types of averages were obtained: those for the individual heads from the four awns of each head; and those for the strain from the averages of the ten heads of each strain. The average values of all of the factors studied, together with their standard deviations and coefficients of variability, were obtained for 1929 and 1950. Differences between.the different factors were obtained by the formula: AiPoEeA ".3 B*PeEeB .. Difference i PeEediffe lO ,Differences were assumed to be significant when the quotient of the difference divided by the probable error of the difference was 5.2 or greater. Correlation coefficients were calculated by the diagonal method used by the Farm Craps Department of Michigan State College which is a modification of the method given by Crum and Patton (2). The limits of significance of r as used in this thesis are: slight - r greater than .3 and greater than six times its probable error. marked - r greater than .5 and greater than six times its probable error. strong - r2 greater than .5 and greater than six times its probable error. RESULTS The data given in Table 1 were obtained by using the average values of each individual head. This table gives the mean, standard deviation, and coefficient of variability with the respective probable errors of each of the five factors studied for the six-row and two-row types of 1929 and 1950. Table 2 contains similar data as Table 1, except that the values were obtained by using the strain averages instead of each individual head. Table 5 gives a comparison of the mean values of each of the factors obtained from individual head averages 11 with those from strain averages. The differences in the values of the means obtained by the two methods are shown to be very slight and in no case is the difference significant. Table 4 gives a comparison of the values of the co- efficients of variability of each of the factors as obtained from individual head averages with those from strain averages. The differences in the values of the co- efficients of variability are large in all cases, the differences being significant for all factors in all of the groups except awn-barbed area ratio and first count six— row 1929 and lengths of barbed area six~row 1950. Table 5 gives a comparison of the means of the five factors for the two-row with those for the six-row strains. The two-row strains have longer awns and longer barbed area than the six-row strains for both years. There was no significant difference between the awn-barbed area ratios in 1929, but in 1950 this value was significantly higher in the two-row strains, showing that they had smoother awns than the six-row strains for that year. The sixprcw strains had a significantly higher "first count" both years. The six-row strains had higher "second count" than.the two-row strains both years, but the differences could not be considered significant. These counts show that the six-row strains had a greater number of barbs per unit area than the 12 two-row’strains. Table 6 gives a comparison of the values of the means of the five factors of the 1929 crOp with those of the 1930 crop, for both two-row and six-row strains. The length of awn was significantly greater in 1930 than in 1929 in.both the six-row and two-row strains. The length of barbed area was greater in 1929 than in 1930, the difference being significant in the two-row strains, but not in the six-row strains. The awn-barbed area ratio was significantly larger in 1930 than in 1929 in'both groups. The "first count" and ”second count" were significantly higher in 1929 than.in 1930 in both groups. This Shows the awns to be smoother in 1930 than in 1929. Table 7 gives a comparison of the values of the co- efficient of variability of the five factors for the two- row’with those for the six-row strains for the two years, 1929 and 1930. In 1929 all of the factors were significant- ly more variable in the six-row than in the two-row varieties, except length of barbed area, which was signif- icantly more variable in the two-row than in the six-row strains for that year. In 1930 the two-row strains were more variable than the six-row strains. This difference was not significant for length of awn but it was for all the other factors. 13 Table 8 gives a comparison of the values of the coefficients of variability of the five factors in 1929 with those in 1930 for the two groups. The length of awn was significantly more variable in 1929 than in 1930. The length of barbed area was slightly more variable in 1929 than in 1930 in the six-row strains. The length of barbed area was significantly more variable in 1930 than in 1929 in the two-row strains. The awn-barbed area ratio in the six-row strains was significantly more variable in 1929 than.in 1930. In the two-row strains the awn-barbed area ratio was more variable in 1930 than in 1929, but the difference was not significant. The "first count" of the six-row'strains was significantly more variable in 1929 than in 1930. The "first count“ of the two-row strains was more variable in 1930 than in 1929, but the difference was not significant. The "second count" of the six-row varieties was more variable in 1929 than in 1930 but the difference was not significant. The ”second count" of the two-row varieties was significantly more variable in 1930 than in 1929. This table shows that the six-row strains were more variable in 1929 than.in 1930, and that the two- row strains except for the length of awn, was more variable in 1930. Whether significant inherited and seasonal differences exist in the degree of smoothness or in the length of awn of l4 two-row strains was determined from a group of 37 strains, all of Which came from the same cross. The mean, standard deviation, and coefficient of variability of four factors were calculated. The several factors of the individual strains for 1929 were compared with the same factors for 1930. Comparisons were made between the values of the mean and the values of the coefficient of variability in all cases. These results are shown in Tables 9 to 16. Table 9 gives the comparison between.the values of the mean of the length of awn for 1929 and those for 1930. Thirtypone of the strains had significantly longer awns in 1930 than.in 1929; four were longer in 1930, but not signif- icantly so; and two were longer in 1929 than.they were in 1930, but these differences were not significant. Table 10 gives the comparison between the values of the coefficient of variability of the length of awn for 1929 and those for 1930. Seven of the strains were more variable in 1930 than in 1929, but none of these differences were significant. Thirty of the strains were more variable in 1929 than in 1930, but in only five cases were the differences significant. Table 11 gives the comparison between the values of the mean of the length of barbed area for 1929 and those for 1930. Seventeen of the strains had longer barbed areas in 1930 than in 1929, but only six of these differences were 15 significant. Twenty of the strains had longer barbed areas in 1929 than in 1930, but only eight of these differences were significant. Table 12 gives the comparison between the values of the coefficient of variability of the length of barbed area for 1929 and those for 1930. Twenty-two of the strains were more variable in 1930 than.in 1929, but in only two cases were the differences significant. Fifteen of the strains were more variable in 1929 than in 1930, but in only two cases were the results significant. Table 13 gives the comparison between the values of the mean of the "first count" for 1929 and those for 1930. In 13 strains the value for the "first count" was higher in 1930 than.in 1929 but in only five cases were these differences significant. In 24 strains the value for the "first count" was greater in 1929 than in 1930; 11 of these strains had significant differences. Table 14 gives the comparison between the values of the coefficient of variability of the "first count" for 1929 and those for 1930. Fourteen of the strains were more variable in 1930 than in 1929; four of these differences were significant. Twenty-three of the strains were more variable in 1929 than in 1930; four of these differences being si gnificant . 16 Table 15 gives the comparison between the values‘ of the mean of the "second count" for 1929 and those for 1950. Twenty-two of the strains had larger "second counts" in 1930 than in 1929; nine of these differences were significant. Fifteen of the strains had larger "second counts” in 1929 than in 1930; eight of these differences being significant. Table 16 gives the comparison between the values of the coefficient of variability of the "second count" for 1929 and those for 1930. Eighteen of the strains were more variable in 1930 than.in 1929, but only four of these differences were significant. Nineteen of the strains were more variable in 1929 than in 1930, but only two of these differences were significant. The possibility of selecting for extreme types of smoothness within these smooth awn segregates was determined by calculating the mean, standard deviation and coefficient of variability, for each of four factors measured, for four of the strains and the three varieties Spartan, Michigan Black Barbless and Michigan Two Row. The four strains used were strains number 70516, 71502, 710206 and 728612. The first two strains are extremely smooth, smooth awn strains, and the latter two are extremely rough, smooth awn strains. {The values of these constants together with the probable error of each, for these four factors are reported in Table 17 17. The values of the constants of these four factors, and the probable errors of each, of the seven varieties and strains, are reported in Table 17. The mean values of length of barbed area and "second count" are shown graph- ically in.Figure 1. Plates 1, 2, 3 and 4 are photo- micrOgraphs of the awns of these four strains. Spartan, Michigan.Two-Row, strains 70516, 71502 and 710206 are two- row; Michigan Black Barbless and strain 728612 are six row. Strains 70516 and 71502 both have significantly shorter awns and barbedareas, and significantly fewer number of barbs than Spartan. Strain 710206 has significantly shorter awns, significantly longer barbed area and a significantly greater number of barbs than Spartan. Strain 728612 has significantly longer awns and barbed area, and a signif- icantly greater number of barbs than Michigan.Black Barbless. Michigan Two-Row, which is a rough awn variety, has signif- icantly longer barbed area than any of these smooth awn varieties and strains, and significantly greater number of barbs than any of these except strain number 728612. Table 18 shows the differences in variability among the five factors for the six-row strains in 1929. Named in order from the least variable to the most variable they arc: length of awn, "first count", length of barbed area, awn- barbed area ratio, "second count". These differences are all significant except the difference between "first count" and 18 length of barbed area. Table 19 shows the differences in variability among the five factors for the six-row strains in 1930. Named in order from the least variable to the most variable they arc: length of awn, "first count", awn-barbed area ratio, length of barbed area, "second count". The differences were all significant. The order of variability was the same for the two years except awn-barbed area ratio and length of barbed area were reversed. Table 20 shows the differences in variability among the five factors for the two-row strains in 1929. Named in order from the least variable to the most variable they arc: length of awn, Ffirst count“, awn-barbed area ratio, length of barbed area, "second count". These differences are all significant between the various factors except the difference between awn-barbed area ratio and length of barbed area. Table 21 shows the differences in variability among the five factors for the two-row strains in 1930. Named in order from the least variable to the most variable they arc: length of awn, ”first count", awn-barbed area ratio, length of barbed area, "second count“. These differences are all significant. The coefficients of variability of the five factors in the two-rOW'group,are in.the same order for the two years, and are in the same order that they are in the l9 six-row strains except for the one difference in the six- row in 1929 which was noted above. The correlations that were computed to ShOW’the relationships between the different factors studied, are reported in Tables 22, 23, 24, and 25. Table 22 contains a report of the correlations that were computed for both the six-row and two-row groups for the two years. The correlations reported in.this table were obtained by using the averages of each individual head. There was a very slight degree of association in all cases between length of awn and length of barbed area, and in 1930 in both the six-row and two-row groups, there was a slight positive correlation between these two factors. There was no correlation between the factors, length of awn and "second count", except in the two-row group for 1930 there was a slight positive correlation between these two factors. There was a strong positive correlation between length of barbed area and ”second count" in all cases. There was a strong negative correlation between “first count" and awn-barbed area ratio in 1929, and a marked correlation between these two factors in 1930. There was a strong negative correlation between "second count“ and awn-barbed area ratio in all the groups.) The latter three correlations show that the greater the number of barbs the longer the barbed area. 20 The same group of correlations were computed using the averages of the strains, instead of each individual head average. This group of correlations is shown in Table 23. The results obtained by this method give very similar results to those obtained by using the averages of each individual head. Comparisons are made between the results obtained by these two methods, these comparisons being shown in Table 24. Only one case shows a significant difference between.the results. This case involves the correlations between "first count" and awn-barbed area ratio for the two-row group in 1929; this difference is 6.83 times the probable error of the difference, although there is a marked negative correlation iriboth cases. Correlations were computed between the factors, length of awn, length of barbed area, awn-barbed area ratio, "first count", and "second count" for 1929 with the same factors for 1930. These are reported in Table 25. There were marked positive correlations between length of barbed areas, between awn-barbed area ratio, between second counts, in both the six-row and tweerow groups and first counts in the six-row group. The other correlation values are not statistically significant. 21 DISCUSSION In making comparisons between the results obtained by using the individual head averages and by using the averages of the strains the values of the means of all the factors were the same from both methods of calculation, ex- cept that the coefficients of variability of all factors obtained by using the individual head averages were higher than the values of this constant obtained by using the strain averages. This is to be expected as some of the strains are quite heterozygous, but the extremes are averaged in to obtain the strain average so that the spread of the pOpulation is very much less than When each head is used to obtain these constants. The results of this experiment show considerable differences between the values of the factors for 1929 When compared with the corresponding values for 1930. A possible explanation is the general difference in the two seasons. The year 1929 was a very poor year for barley and the plots used in 1929 were very low in fertility. As a result the yields were so low in 1929 that only enough of each strain was harvested to obtain seed for the 1930 planting. Differences in all the factors studied were found to exist between the six-row and the two-row groups. These differences, while due to differences in inheritance, are 22 modified by seasonal conditions. The two-row strains had significantly longer awns than the six-row strains, but this difference was not so great in 1930 When'better grow- ing conditions prevailed. The six-row strains had the shorter barbed area both years but this difference was less in 1930. There was no difference in the awn-barbed area ratio in 1929, but in 1930 this factor was significantly larger for the two-row group than it was in the six-row group. The "first count” was significantly less for the two-row than it was for the six-row both years, but the difference was greater in 1950 than it was in 1929. The "second count" was less for the two-row strains both years but the differences were not significant either year. These results show'that the two-row strains have longer and smoother awns than.the six-row strains, but that the differences vary with seasonal conditions. The length of awn in 1930 was significantly greater than in 1929 in both the six-row and two-row groups but the coefficient of variability was low both years. This shows that there are differences due to seasonal conditions and that differences due to inheritance were slight. The awn- barbed area ratio was greater and the number of barbs was lower in 1930 than in 1929. The coefficients of variability in these 12 dases were high showing that there were differences in smoothness of awn which were caused by differences in inheritance and by differences in seasonal conditions. A group of the two-row type strains were used to ' make comparisons between the values of four factors studied in 1929 with their corresponding factors in 1930. The results of this study correspond very closely to the results obtained by comparing the entire pOpulation of 1929 with that of 1950. A comparison was made between.hfichigan Black Barb- less and Spartan which are smooth awn.varieties and Michigan Two-Row Which is a rough awn variety wdth two extremely rough awn strains and two extremely smooth awn strains. The results of these comparisons ShOW’that all of the strains are much smoother than Michigan Two-Row, but that there are wide differences between strains that are "smooth awn" selections from smooth awn x rough awn crosses. Thesecomparisons also show that it is possible to obtain strains that are smoother than the present commercial smooth awn.varieties. In making comparisons among the different factors to determine the amount of variability it was found that they rank in the following order: length of awn, "first count", awn-barbed area ratio, length of barbed area, and "second count", named in order from.the least variable to the most variable. All of the strains have a number of 24 barbs near the tip but there is a wide variation between strains in the distance they extend down the awn, Which ' fact is brought out by the results. The results of the correlation studies show that smooth awn is not associated with length of awn, and that the long awned strains are as likely to be smooth as are the short awned strains. There is a very close association, however, between the length of the barbed area and the number of barbs "second count". Some strains had very few barbs per unit area but they extended quite far down the awn Which tends to lower this correlation. This fact also tends to make the correlation between awn-barbed area. ratio and "first count" lower than the correlation between awn-barbed area ratio and "second.count". The relatively high negative correlation between awn-barbed area ratio with both "first count" and "second count" show that, if the barb counts are high, a large portion of the awn will be barbed. The inter-annual correlations of all of the factors for both six-row and two-row strains were positive and significant, except for length of awn and "first count" in the two-row group and length of awn in thesix-row group. 'The high positive correlation between length of barbed areas between awn-barbed area ratios, between "first counts and between "second counts" show'that these factors are 25 quite constant within a strain and not so dependent up- on the environment as length of awn. SUMMARY 1. It is as reliable to use the strain averages as the individual head averages in computing the mean of any one of the factors, but the individual head averages should be used to obtain the coefficient of variability. 2. There are significant differences in smoothness of awn and length of awn due to inherited and seasonal differences. The two-row group had longer and smoother awns than the six-row group both years. Both of the groups had longer and smoother awns in 1930 than in 1929. 3. There are significant differences in degree of smoothness and length of awn among the strains in the six- row group Which are due to inherited and seasonal dif- ferences. 4. There are significant differences in degree of smoothness and length of awn among the strains in the two- row group Which are due to inherited and seasonal dif- ferences. 5. There are significant differences in length of awn among a group of two-row strains due to inherited and seasonal differences. There also were significant dif- A 26 ferences in smoothness of awn among some of these strains due to seasonal and inherited differences, but the numbers used were probably too small to bring out all of the dif- ferences. 6. There are "smooth awn" strains that are smoother and "smooth awn" strains that are rougher than Michigan Black Barbless and Spartan. 7. Length of awn was the least variable factor studied, followed in order by "first count", awn-barbed area ratio, length of barbed area, and "second count". This shows that all of the strains had a number of barbs near the tip, but there was a wide range of variation in the distance they extended down the awn. 8. There was a strong correlation between number of barbs and length of barbed area, and between number of barbs and awn-barbed-area ratio. The correlation between length of awn and length of barbed area and between length of awn and number of barbs was very low in all cases. In making selections smooth awn strains could be selected by determining the distance the barbs extended down the awn. There is a strong degree of association between this factor and number of barbs, so the awn would be relatively smooth if the barbs did not extend over more than a third of the awn. All of the factors for smoothness of awn for one year showed a high degree of association with the same 27 factor for the next year. This shows that these factors were quite constant within a strain and so inherited. LITERATURE CITED 1. Colin, H. and Trouard-Riolle. Dissociation of the Barley Hybrid Smooth Awned Black x Rough Awn.Albert; Int. R of Sci. and Prac. Agr. N. S. 1:655. 1925. 2. Crum, W. L., Patton, A. C. An Introduction to the Methods of Economic Statistics. New‘York. A. W. Shaw Co. 1925. 5. Griffee, Fred. Correlation Inheritance of Botanical Characters in Barley and manner of Reaction to Helminthosporium Sativum; Jour. of Agr. Res. 50: 915-955. 1925. 4. Harlan, H. V., Hayes, H. K. Investigations in Barley Breeding. Minn. Agr. Exp. Station Bul. 182: 45-56. 1919. 5. Smooth Awned Barleys. Jour. Am. Soc. of Agron. 12:205-228. 1920. e. and Pope, M. N. The Use and Value of Back Crossing in Small Grain Breeding. Jour. of Heredity 15: 519-522. 1922. 7. Hayes, H. K. and Wilcox, A. M. The Physiological Value of Smooth Awned Barleys. Jour. Am. Soc. Agron. 14: 113. 1922. 8g 9. 10. 11. 12. 28 Hayes, H. K. Breeding Improved Varieties of Smooth Awned Barleys. Jour. of Heredity 17: 571-581. 1926. Hor, K. S. Interrelation of Genetic Factors in Barley. Genetics 9(2): 151-180. 1924. Rather, H. 0., Down, E. E., Brown, H. M., Clark, F. H. Barley for Michigan Farms, Michigan Agricultural Experiment Station Special Bulletin 191, march 1929. Sigfusson, S. J. Correlated Inheritance of glume color, Barbing of Awns, and Length of Rachilla Hair in Barley. Sci. Agr. 9:662-674. 1929. Vavilon, H. 0. Origin and Genetic Composition of Types of Smooth Awned Barley. Int. R Sci. and Prac. Agr. N. S. 1, p 952. 1925. Table 1. The Means, standard deviations, and coeffi- cients of variability with their Probable Errors of the five factors studied, using the averages of each individual head for the two-row and six-row types in 1929 and 1950. Standard Coefficient Mean Deviation of Var. Length of awn six-row, 1929 111.25£.58 14.81t.27 15.5lt.25 six-row, 1950 125.75i.29 ll.97£.21 9.52t.l6 two-row, 1929 l52.41¢.50 15.70*.21 11.86i.l6 two-row, 1950 140.851.24 15.50i.17 9.591.12 Length of barbed area six-row, 1929 48.55t.40 15.46:.28 51.99t.64 Six-row, 1950 47.611.55 14.421.25 50.29%.57 two-row, 1929 58.l5¢.58 20.541.27 55.14¢.52 two-row, 1950 55.591.42 25.441.29 45.891.65 Awn-barbed area ratio six-row, 1929 2.57i.05 1.011.02 59.161.81 81XP1‘0W, 1930 2.841002 onteOI 24.94te46 two-row, 1929 2.54t.02 .851.0l 55.59:.49 tIVO'TOW, 1930 3008*002 10091.01 55.47‘050 First count six-row, 1929 68.59£.55 21.611.59 51.51£.65 six-row, 1950 66.40:.57 15.081.26 22.72:.41 two-row, 1929 64.25:.52 16.911.22 26.55t.57 two-row, 1950 60.591.29 16.501.20 27.001.56 Second count SiX'I‘OW, 1929 55o57to48 18058i035 52.251018 six-row, 1950 52.24:.57 15.551.27 47.551.99 two-row, 1929 55.961.50 15.771.21 46.441.74 two-row, 1950 50.971.52 17.91i.22 57.84:.80 TablO 20 Same determinations as in Table 1, except computed from.averages of each strain. Mean 'Length of awn SiX’rOW, 1929 111059£088 SiX'rOW, 1930 125081*070 two-row, 1929 151.881.65 tWO'TOW, 1950 1400741063 Length of barbed area six-row, 1929 48.21t.98 SiX’IOW, 1950 47076t097 two-row, 1929 58.501.95 two-row, 1950 55.45t1.15 Awn-barbed area ratio SiX'rOW, 1929 2051*007 six-row, 1950 2.80:.04 two-row, 1929 2.46t.04 two-row, 1950 2.99i.05 First count six-row, 1929 68.1511.45 six-row, 1950 65.752.95 two-row, 1929 64.161.79 two-row, 1950 60.221.75 Second count six-row, 1929 55.05£1.16 SiX‘IOW, 1950 31059$090 two-row, 1929 55.811.65 two-row, 1950 50.98:.85 Standard Deviation ll.50t.65 9.141.50 11.26:.45 11.26:.45 12.761.69 12.64i.69 16.951.67 20.211.80 .86i.05 .571.05 0662003 .92t.04 18.6611.0l 12.551.67 14.031.56 15.50:.55 15.111.82 11.70:.64 11.67i.46 14.80t.59 Coefficient of Variability 10.14i.46 7.261.39 8.541.34 8.001.22 26.46i1.54 26.46t1.54 25.49i1.07 57.82i1.70 54.59t2.08 20.16t1.14 26.8111.13 50.8611.53 27.58tl.60 18.78t1.06 21.8710.91 22.08i0.92 45.13i2.74 37.05t2.59 34.52t1.52 47.7711.08 q Table 5. Comparison between the value of the means re- ported in Table 1 and the corresponding value reported in Table 2. Individual head Strain Difference Difference 0 ‘00 l 0 Length of awn six-row, 1929 111.25i.58 111.59i.88 .561.96 0.4 six-row, 1950 125.75t.29 125.81i.70 .06t.76 0.1 two-row, 1929 ‘l52.41t.50 151.881.65 .55£.70 (3.8 two-row, 1950 140.851.24 140.74t.65 .091.67 0.1 Length of barbed area six-row, 1929 480332040 480211’098 01211006 001 six-row, 1950 47.611.55 47.761.97 .1511.05 0.2 two-row, 1929 58.15:.58 58.401.95 .57.+1.02 0.4 tWO"I'0W, 1930 530391042 530431'1015 00411021 000 Awn-barbed area ratio SiX‘rOW, 1929 2057t005 20511007 006*008 O08 SiXpl‘OW, 1950 2089*002 2080*004: 009*004 203 two-row, 1929 2.541.02 2.462.04 .08i.04 2.0 two-row, 1950 5.081.02 2.9%.05 .091.05 1.8 First count 81X'1‘0W, 1929 68059t055 6801531045 0441'1055 003 SiX’rOW, 1930 66040t057 65.75’095 06511002 006 two-row, 1929 64.251.52 64.162.79 .091.85 0.0 two-row, 1950 60.593.29 60.22:.75 .1fli.80 0.0 Second count six-row, 1929 55.571.48 55.05t1.l6 .5411.26 0.4 six-row, 1950 52.241.57 51.591.90 .65:.97 0.7 tm’row, 1929 35096‘030 330816065 015:072 002 tWO'I‘OW, 1950 300971052 30098i085 .OM.89 000 Comparison of the values of the coefficients of variability reported in Table l, and the corres- ponding values reported in Table 2. Table 4. Individual head Length of awn six-row, 1929 15.51:.25 six-row, 1950 9.52:.16 two—row, 1929 11.861.16 tWO'IOW, 1950 90593012 Length of barbed area 81X‘rOW, 1929 310991064 six-row, 1950 50.29:.57 two-row, 1929 55.14:.52 two-row, 1950 45.89:.65 Awn-barbed area ratio six-row, 1929 59.161.81 six-row, 1950 24.94:.46 two-row, 1929 55.59t.49 two-row, 1950 55.47i.50 First count six-row, 1929 51.51t.65 six-row, 1950 22.721.41 two-row, 1929 26.551.57 two-row, 1950 27.00:.56 Second count six-row, 1929 52.2511.18 six-row, 1950 47.551.99 two-row, 1929 46.44:.74 two-row, 1950 57.841.80 Strain 10.14£.46 7.261.59 8.54%.54 8.001.22 26.46£1.54 26.46t1.54 26.49¢1.07 57.82£1.70 54.39i2.08 20.16t1.14 26.81t1.15 50.86i1.33 27.5811.60 18.7811.06 21.87:.91 22.083.92 43.13i2.74 37.0312.59 54.5211.52 47.7711.08 Difference Difference 5.171.52 2.261.42 5.524.58 1.59*.25 5.5511.67 3.8311.64 8.65:1.19 6.0711.82 4.7712.23 4.78i1.25 6.58t1.23 V4.61t1.42 4.13:1.72 5.94i1.14 4.469.98 4.92:.99 9.12i2.98 10.50t2059 1109221069 1000721034 P.E.o 6.1 qqfu mHHH Di . Table 5. Comparison between the value of the means of the ‘ five factors studied for the six-row group, and the corresponding means of the two-row group. Length of awn Length of awn Length of barbed area Length of barbed area Awn-barbed area ratio Awn-barbed area ratio First count First count Second count Second count Six-row lurges 1929 1930 1929 1930 1929 1930 1929 1950 1929 1950 + U - Six-row smallest Six-row 111.23£.38 125.751.29 48.33to40 470611035 2.571.05 2.841.02 68.591.55 66.401.37 55.57i.48 52024t057 Two-row 58.13t.38 53.39i.42 2.541.02 5.081.02 64.25t.32 60.591.29 33096‘050 30.971.32 Difference Difference 152.411.50 -21.18i.48 140085t024 “150081038 9.801.55 5.78t.55 .05t.04 .24:.05 4.341.64 6.01:.47 1.61i.57 1.271.49 P.E.Diff. 44.1 39.7 17.8 10.5 0. 8. 6. 12. 2. 2. Oflb END CHD Table 6. Comparison between the value of the means of the five factors studied for 1929 and the corresponding means in 1950. 1929 1950 Difference Difference P.E.DiII. Length of awn SiX'I‘OW 1110231’058 125075i029 "140521048 3003 Two-row 132.4lt.50 140.83t.24 - 8.421.58 22.2 Length of barbed area Six-TOW 48035i040 470 61*0 35 072*053 104 Two-row 58.131.38 53.391.42 4.74t.57 8.3 Awn-barbed area ratio Six-row 2.57f.03 2.841.02 - .27t.04 6.8 TWO’I‘OW 2054*002 30081002 _ 0541003 1800 First count Six-row 68.59t.55 66.401.37 2.19£.66 3.3 TWO'I'OW “025:032 60039i029 30861043 900 Second count SiX‘I‘OW 350571048 320241037 3033*0 61 505 TWO‘I'OW 530961050 300971032 2099:044 608 - 1929 smallest Table 7. Comparison between the values of coefficients ‘ of variability of the five factors studied for the six-row group and the corresponding values for the two-row group. Six-row Two-row Difference Difference P.E.Diff. Length of awn 1929 15.51£.25 11.864.16 1.45:.50 4.8 Length of awn 1930 9.521.16 9.59t.12 - .07t.20 0.4 Length of barbed area 1929 51.99£.64 55.141.52 -5.15t.82 5.8 Length of barbed area 1930 50.291.57 45.891.65-15.601.86 15.8 Awn-barbed area ratio 1929 59.16t.81 53.59:.49 5.771.95 6.1 Awn-barbed area ratio 1950 24.94:.46 55.471.50-10.531.68 15.5 1 First count 1929 51.511.65 26.55i.57 5.18:.75 7. First count 1950 22.72t.41 27.00£.56 -4.284.55 7.8 Second count 1929 52.25il.18‘46.44i.74 5.811l.59 4.2 Second count 1930 470551099 57084*080‘10031*1027 801 - Six-row smallest Table 8. Comparison between the values of the coefficients of variability of the five factors studied in 1929 and the corresponding values for 1950. 1929 1930 Difference Difference F.E.Diff. Length of awn SiX'rOW 130:31t025 9052i016 3079*030 1206 two-row 11.861.16 9.59t.l2 2.271.20 11.4 Length of barbed area six-row 51.991.64 50.29£.57 1.601.86 2.0 two-row 55.141.52 45.891.65 -8.75t.83 10.5 Awn—barbed area ratio SiX‘I‘OW 39016t081 24094’046 l4022§093 1503 two-row 53.39:.49 55.47:.50 -1.98t.70 2. First count six-row 51.519.65 22.72t.41 8.791.75 11.7 two-row 260551057 270001056 " 0671052 103 Second count six-row 52.25t1.18 47.53t.99 4.72:1.54 5.0 two-row 46.44:.74 57.841.80 -ll.4011.09 10.5 -- 19:32) 3 allest Table 90 1929 152.031lo68 130.48t2.85 118.0802.38 118.9331.92 124.39t1.68 127.25¢2.12 119.38t2.12 120.08t1.60 124.7745.58 111.4813.3O 150.3112.25 123.39i4.03 135.95f3.14 145.10t2.60 134.4 (34.37 143.09t2.19 127.05t2.57 147.38i3.15 159.0543.54 136009f5002 152.73£1.68 135.42t4.70 130.3813.58 144.5012.45 138.7713.23 137.4712.96 146.1552.71 123.3412.89 124.8512.18 l24.18£3.06 126.3512.93 118.14i3.23 125.00t2.78 154.4512.56 136.36f2.55 120.00*1.80 127.9512.17 1930 152.98t2.62 142.931.78 127.23tl.36 134.4811.45 139.3502.01 136.801.80 137.48i1.77 146.2311.68 158.8012.08 134.5544.31 163.10t2.31 141.00t6.67 150.00il.59 149.9811.55 150.08t2.20 152.60tlo58 l46.80£2.27 146.15f1.82 152.08i3.40 150.85t1.86 157.70i1.46 161.08i3.04 158.2542.79 141.0341.72 155.60i2.38 157.43é1.44 166.75él.78 149.18i2.11 l52.50£3.23 148.55t2.09 l65.75$1.30 l39.9011.94 l54.50¢1.79 161.33i1.11 162.33t1.78 140.95t3.05 145.1311.05 1950 largest - 1930 smallest Difference .95i5.ll 12.4512.95 9.15t2.74 l5.58t2.40 14.96t2.62 9.55f2.27 18.10i2.76 26.15i2.32 54.0313.97 25.07i5.43 32.7943.22 17.6127.79 14.0543.52 4.8843.05 l5.68$4.89 9.51t2.70 19.7545.45 ‘ 1023i3064 l5.03t4.77 14.7613.54 4097*2023 27.6645.60 27.87i4.54 - 3.4712.99 16.83t4.01 l9.9613.29 20.60t5,24 25.84t3.54 27.6543.90 24.3713.7l 59.4043.20 21.7643.77 29.50:5.31 26.8802.79 25.9715.ll 20.9545.54 17.1842.41 Mean lengths of awn, 1929 and 1950, of a group of two-row strains. Difference 0.40 0 .3 H qwmommmmqqmmmt—‘mpmsm (numl-‘IPNOIPCDHOIOPUIO‘INP [—1 OOOOOOOOOOOOOOOOO .00... O H‘DIPOW‘OCDNOHNPHNNHQNNfiWQCfiNO‘ONNCfiO‘CflO‘NdU‘CflN Table 100 1929 5.18i.78 10.21tl.55 9.4521.44 7.20ilol5 5.28*.95 7040*1017 805111.27 6.23*.94 12.04tlo94 13.1702.13 7.6801.25 14052£2023 10082*1065 804111027 13063§2034 6042£1008 9049*1043 10.03*l.53 8.74el.70 902921057 5.17$.7B 12078*1051 12.88i1098 7095f1020 10.3411.66 900421052 8071*1.51 9084*1067 8.18i1.25 11.55t1077 10088*1066 12.83i1097 10.43t1073 8.9111.34 8.5131032 6068*1006 7094*1020 1930 8004*1022 2.554.58 40991.76 5.071.76 607721003 2.741.41 6.049.91 5.574.81 6.16t.93 17.22i2.67 6.6511.00 21.02t3.30 4.95:.75 4085;075 6.8811.04 4.85:.75 702411009 5.84t.88 10.84*1.6O 5.744.87 4.33i.67 8.85$1.33 8.27t1.25 5.730.86 7.17*1.08 4.28:.65 5.01:.76 606331000 9.86!1.5 60581099 5067t055 6.50:.98 5.42:.82 5.22:.48 5.151.78 10013*1054 3038*051 1950 largest - 1950 smallest Difference 208611044 “706611060 “404611065 “201321038 1049t1040 “406611024 “202731056 ' .8611.24 -5.88*2.15 400513042 “100331059 6.50:5.98 “5087*1081 -5.56tl.47 “6075f2056 “1057*1030 -2.25tl.80 '4.19tl.77 2010*2033 “505511080 - .84tl.03 '3093£2.Ol “4061*2034 “2022*1048 “5017$1098 “4.76t1.65 “5070*1051 “5021*1094 .684l.95 “409812003 -7.2111.75 “6033*2020 “5.0131096 “5069*1042 “501651053 5.45tl.87 ‘4.56$1.50 Coefficients of variability of the length of awn, 1929 and 1950, of a group of two- row strains. Difference 0 0 l 0 .0... OOOOOOOOOOOOOOO. (flFJNHPtODDFKO FHMDDFHJDDN> D0 IDFJFWDRDUH4 F‘Ni F‘UH‘PJNHPIO 0 UHDF‘CDOMOP‘UHF-QUHOCDUHDCDGHDQDQNNDOOHF[OOVQtO