Euphytica 33 (1984) 907-913

INHERITANCE OF GROAT PROTEINPERCENTAGE IN AVENA SATIVA L . x A . FATUA

L . CROSSES

J . M. REICH and M . A. BRINKMAN

Department of Agronomy, University of Wisconsin, Madison, WI 53706, USA

Received 17 April 1984

INDEX WORDS

Avena saliva L ., oat, Avena fatua L., wild oat, interspecific oat crosses, groat protein percentage, inheritance,seed shattering .

SUMMARY

Gene action and heritability of groat protein percentage were determined in F,, F 2 , and F 3 generationsof nine crosses between three A vena saliva L . cultivars and three A. fatua L . selections . Relationships amonggroat protein percentage, grain yield, and 100-seed weight also were evaluated . The three A. saliva parentswere Dal (high grain yield and intermediate groat protein percentage), Goodland (low grain yield andhigh groat protein percentage), and Stout (high grain yield and low groat protein percentage) . The threeA. fatua parents were chosen for the study on the basis of vigorous plant growth and high groat proteinpercentage . The study was conducted at Madison, Wisconsin in 1979 and 1980 .

There was partial dominance towards low groat protein percentage . Narrow sense heritability estimatesfor groat protein percentage were low in Dal and Goodland crosses and intermediate in Stout crosses .In the F 2 generation, groat protein percentage was significantly higher in shattering than in nonshatteringplants in 1979, but not in 1980. There were significant, positive correlations between groat protein percent-age, 100-seed weight, and grain yield in F, and F 2 generations, but they were not large numerically . Relation-ships among these traits were either negative or nonsignificant in the F 3 generation . Although our resultsindicated that selection for higher groat protein percentage is possible when a low protein A. saliva cultivaris used, most of our simple cross progenies from A. saliva x A . fatua crosses had weak straw and weresusceptible to crown rust (Puccinia coronata CDA . var. avenae FRASER and LED .) .

INTRODUCTION

Introducing exotic germplasm into cultivated oat (Avena sativa L.) breeding programshas been common in recent years . Of the two prevalent hexaploid wild oat species,the wild red oat (A . sterilis L .) has been used more extensively than the wild oat (A .fatua L.) . Resistance to crown rust (Puccinia coronata CDA . var . avenae FRASER andLED.) has been the most widely coveted characteristic in A. sterilis, although severalstudies have reported improvements in agronomic and quality characteristics incrosses with this species (FREY & BROWNING, 1971 ; CAMPBELL & FREY, 1972a ; FREY,1976; TAKEDA & FREY, 1979) . Collections ofAafatua have not been used as extensivelybecause they do not have as much resistance to crown rust as A. sterilis (BRIGGLEet al., 1975 ; RINES et al ., 1980) . Nevertheless, A. fatua is characterized by vigorous,

907

J . M . REICH AND M . A. BRINKMAN

Table 1 . Pedigrees of A. sativa cultivars and descriptions of A. fatua collections used as parents in thestudy.

competitive growth and groat protein percentages that are higher than A. sativa (RINEset al ., 1980) .

There is evidence that groat protein percentage in A. sativa x A . sterilis crossesis under genetic control . Several studies have indicated significant additive gene actionfor this trait, with partial dominance towards low protein percentage in early genera-tions (CAMPBELL & FREY, 1972b ; SPILDE et al ., 1974; SRAON et al., 1975 ; IWIG & OHM,1976). Heritability estimates in these studies have ranged from 9 to 76%, with mostestimates in the 40 to 50% range .

Virtually all of the studies on groat protein percentage and its relationship to otheragronomic traits in adapted x exotic oat crosses have dealt with A. sativa x A. sterilisprogenies. The objective of this study was to evaluate A. sativa x A . fatua crossesto determine (a) heritability and gene action for groat protein percentage, and (b)relationships among groat protein percentage, grain yield and 100-seed weight .

MATERIALS AND METHODS

Three A. fatua selections were used as male parents in all possible crosses with A .sativa cultivars Dal (high grain yield and intermediate groat protein percentage),Goodland (low grain yield and high groat protein percentage) and Stout (high grainyield and low groat protein percentage) (Tables 1 and 2) . The three A. fatua parentswere chosen for the study on the basis of vigorous plant growth and high groat proteinpercentage. Grain yield, groat protein percentage, and 100-seed weight were obtainedfrom individual parent, F 1 , and F2 plants in both 1979 and 1980 and from F3 rowsin 1980. Six to 12 parent and F 1 plants per cross were grown each year . Ninety F 2plants were grown per cross in 1979, while 51 to 66 F 2 plants were grown per crossin 1980 . Eighty F 3 lines derived from randomly selected F 2 plants were grown in eachcross in 1980 .

The study was grown at Madison, Wisconsin, on a silt loam soil that was fertilizedwith 20 kg N, 81 kg P205i and 81 kg K20 per ha. The experimental design was a908

Euphytica 33 (1984)

A . sativa

cultivar

DalGoodlandStout

originating state

WisconsinWisconsinIndiana

pedigree

Trispernia/Belar/2/BeedeeGoodfield/2/Trispernia/Belar/3/Goodfield/4/GarlandPedigree includes Milford, Ukraine, Clinton3 , and Clintland°

A.fatua

collection no . location of collection lemma color pubescence

175 Southern Red River Valley dark brown intermediate955 Oat field, Webster, S .D . white intermediate971 Flax field, Millbank, S.D . grey short

GROAT PROTEIN IN AVENA

Table 2 . Grain yields and groat protein percentages of A . sativa cultivars grown in 1 .5 m2 plots at Madison,Wisconsin in 1976, 1977, and 1978 .

* Within colums, means followed by the same letter are not different at the 0 .05 level of significance accord-ing to DNMRT .

randomized complete block with three replicates in 1979 and two in 1980 . Plantingdates were May 10, 1979 and May 3, 1980 . Adequate soil moisture was presentthroughout 1979, but in 1980 all of May and the first two weeks of June were considera-bly drier than normal in southern Wisconsin (REICH, 1981) . Four applications of Man-zate (zinc manganese ethylene bisdithiocarbamate) were made at weekly intervals afteranthesis to control crown rust .

Except for the F3 rows, the materials in the study were planted in semi-spaced condi-tions. This was essential in order to harvest shattering and non-shattering plants indivi-dually. Row spacing was 0 .3 m. Parents, F,, and F 2 seeds were planted 7 .5 cm apartin rows that were 1 .5 m long (44 plants/m 2 ). F3 lines were seeded in 1 .5 m rows ata rate of 100 seeds/m 2 , which is somewhat less than the normal rate of 150-175 seeds/m2 used in the region when alfalfa (Medicago sativa L .) is being established .

At maturity, individual parent, F,, and F 2 plants were harvested and threshed, andgrain yield, groat protein percentage, and 100-seed weight were determined . The F3lines were harvested as individual rows, and after grain yield and 100-seed weightwere determined, a 5 g aliquot of seed was used to measure groat protein percentage .Samples were prepared for protein analysis by dehulling with an impact-type dehuller,and crude protein percentage was calculated by multiplying Kjeldahl nitrogen percent-age by 6 .25 .

Panicles of shattering plants, identified at anthesis by two long awns protrudingfrom a spikelet, were enclosed in gas permeable cloth bags and staked upright to pre-vent seed loss . Shattering type panicles were bagged and staked approximately twoweeks after anthesis . In each cross, groat protein percentage comparisons betweenshattering and non-shattering classes of F2 plants were made with a t-test (STEEL &TORRIE, 1980) .

Narrow sense heritability for groat protein percentage selection on an individualplant basis was estimated from the linear regression of F 3 progeny rows on F2 parentalplants. Gene action, determined as the degree of dominance, was calculated as de-scribed by PETR & FREY (1966) :

Degree of dominance (F,) = h, =F, - M .P .H.P . - M.P .

where F, M.P ., and H.P. represent the mean F,, mid-parent, and high parent groatprotein percentages, respectively .

Euphytica 33 (1984) 909

Cultivar Grain yield Groat protein(kg/ha) (%)

Dal 2810a* 18 .8bGoodland 2443b 20.3aStout 2750a 17 .3c

Table 3 . Grain yields, 100-seed weights, and groat protein percentages of A. sativa and A. fatua parentsgrown at Madison, Wisconsin in 1979 and 1980 .

J . M . REICH AND M. A. BRINKMAN

Avena fatua parents averaged 21 .6% groat protein and 2.5 g grain yield per plant,while A . sativa parents averaged 17.9% groat protein and 4.7 g grain per plant (Table3). Grain yields averaged over the two years were relatively low because the dry condi-tions in 1980 reduced tiller and spikelet numbers . Grain yields were average in 1979 .

The correlation between groat protein percentage and grain yield among the six par-ents was significantly negative (r = -0.70*) .

Seed characteristics of A . sativa were more desirable than those of A. fatua . Forexample, A. sativa parents had mean 100-seed weights that were nearly twice as largeas the A. fatua parental means (Table 3) . The negative correlation between kernelweight and groat protein percentage was highly significant (r = -0.91 **) .

Dominance values for groat protein percentage were moderately large and negative(Table 4) . Among A. fatua parents, crosses involving collection 955 had the largest

Table 4 . Degree of dominance and narrow sense heritability estimates for groat protein percentage in nineinterspecific oat crosses grown at Madison, Wisconsin in 1979 and 1980 .

910 Euphytica 33 (1984)

Selection or cultivar Grain yield 100-seed Groat protein(g/plant) weight

(g)(%)

A. fatua 175 2 .3 1 .62 22.3A. fatua 955 2 .4 1 .65 20.0A. fatua 971 2 .7 1 .80 22 .5

Mean ofAafatua parents 2 .5 1 .69 21 .6

Dal 4 .9 2 .88 17 .3Goodland 4.4 3 .09 20.3Stout 4.8 3 .43 16.3

Mean of A. sativa parents 4.7 3 .13 17 .9

RESULTS AND DISCUSSION

Pedigree Cross Degree of dominance Narrow sensenumber heritability

1979 1980

Dal/175 1 - 1 .58 - 1 .55 0.16±0 .11Dal/955 2 - 2 .47 - 3 .39 0.13±0 .16Dal/971 3 - 1 .31 - 1 .62 0.06±0 .13

Goodland/175 4 - 5.14 - 3 .27 0.05±0 .07Goodland/955 5 -24.80 -53.00 0.06±0 .08Goodland/971 6 - 4.87 - 4.18 0.12±0 .06

Stout/175 7 - 1 .77 - 0 .55 0.48±0 .11Stout/955 8 - 2 .71 - 0 .88 0.27±0 .13Stout/971 9 - 1 .55 - .00 0.34±0 .09

GROAT PROTEIN IN AVENA

Table 5 . Comparison of groat protein percentages for shattering (Sh) and non-shattering (NSh) classesof F Z plants in nine interspecific oat crosses grown at Madison, Wisconsin in 1979 and 1980 .

*, ** Significant at the 0 .05 and 0.01 levels, respectively .

negative dominance values, while among A. sativa parents those involving Goodlandwere most negative . The large negative dominance values observed for progenies ofGoodland x A . fatua 955 suggest that epistatic gene action may influence groat pro-tein percentage in this cross . Most of the dominance estimates in this study are similarin magnitude to those obtained from A . sativa x A . sterilis crosses (CAMPBELL & FREY,1972b ; OHM & PATTERSON, 1973 ; SPILDE et al., 1974; SRAON et al ., 1975 ; IWIG & OHM,1976) and indicate partial dominance towards low protein percentage .

Narrow sense heritabilities for groat protein percentage, estimated from the regres-sion of F 3 and F 2 , were low for Dal and Goodland crosses and intermediate for Stoutcrosses (Table 4) . Dal and Goodland were intermediate and high protein parents, re-spectively, and Stout was a low protein parent. Crosses with Goodland and Stouthad the lowest and highest heritabilities, respectively, which is not surprising becauseparental differences in groat protein percentage were small for Goodland versus A .fatua but were much larger for Stout versus A . fatua . These tendencies indicate thatselection for higher groat protein percentage in interspecific crosses with A . fatua maybe effective if the A . sativa parent has low groat protein percentage, but selection prob-Euphytica 33 (1984) 911

Year CrossNo.

Shattering F Z plants Non-shattering F Z plants Comparison of shatteringand non-shattering classes

numberofplants x SX

numberofplants x SX

_XSh-NSh S d t

1979 1 21 19 .1 0 .57 69 15 .3 0.18 3 .8 0 .60 6.37**2 21 16 .6 0 .35 69 14 .4 0.15 2.2 0 .40 5.57**3 23 18 .3 0 .44 67 15 .3 0.25 3 .0 0 .50 5 .95**4 23 19 .6 0 .66 67 16 .2 0.27 3 .4 0 .71 4.76**5 21 17 .6 0 .66 69 15 .6 0.23 2.0 0 .70 2.87**6 23 18 .8 0 .72 67 16 .8 0.35 2.0 0 .80 2.517 21 17 .1 0 .67 69 14 .6 0.18 2.5 0 .69 3 .61 **8 22 15 .4 0 .42 68 13 .9 0.11 1 .5 0 .43 3 .46**9 22 17 .2 0 .45 68 14 .8 0.17 2.4 0 .48 5 .00**

Mean 17 .7 15 .2 2.5

1980 1 14 19 .5 0 .56 39 18 .0 0.38 1 .5 0 .68 2.212 17 19 .4 0 .42 43 18 .5 0.20 0.9 0 .46 1 .943 17 18 .7 0 .31 43 18 .3 0 .30 0.4 0 .44 0.924 13 22 .0 0 .53 50 20 .0 0.24 1 .8 0 .53 3 .41**5 15 19 .0 0 .39 51 18 .4 0.35 0.6 0 .53 1 .146 18 20 .2 0 .41 44 20 .2 0.48 0.0 0 .63 0.007 11 18 .3 0 .35 51 18 .3 0.23 0.0 0 .42 0.008 8 17 .0 0 .63 43 17 .6 0.25 0.6 0 .68 0.889 16 18 .1 0 .50 45 18 .6 0.28 0.5 0 .58 0.87

Mean 19 .1 18 .7 0.5

*,** Significant at the 0 .05 and 0 .01 levels, respectively .

ably will be ineffective if the A . saliva parent is intermediate to high in groat proteinpercentage .

Differences in groat protein percentage between shattering and non-shatteringclasses of F 2 plants were influenced by environment . Groat protein percentages aver-aged 2 .5% higher in shattering F 2 plants in 1979, but shattering plants were significant-ly higher in only one cross in 1980 (Table 5) . The 1979 results support the contentionthat shattering per se causes increased groat protein percentage (CAMPBELL & FREY,1972a ; LYRENE & SHANDS, 1974). When this occurs, reduced carbohydrate content(deposition) due to shattering results in increased protein percentage without an in-crease in protein content (LYRENE & SHANDS, 1974) . It is possible that shattering andnon-shattering types did not differ in groat protein percentage in 1980 because thedry conditions in May and June, which reduced grain yields (REICH, 1981), resultedin higher groat protein percentages . With below-average grain yields, groat proteinpercentages of the non-shattering types probably were higher than they ordinarilywould have been. In both years, non-shattering and shattering plants in F2 populationsfit a 3 : 1 segregation ratio .

Although groat protein percentage was positively correlated with grain yield of F 1and F 2 plants, it was negatively correlated with grain yield of F 3 lines which weregrown under more competitive conditions (Table 6) . Four of the five groat proteinpercentage-grain yield correlations in Table 6 were significant, but none of them werelarge numerically. The negative correlation between groat protein percentage andgrain yield of F 3 lines suggests that improving both traits simultaneously would bedifficult. Significant correlations between groat protein percentage and 100-seedweight also were small numerically (Table 6) .

In summary, simple cross progenies from A. saliva x A . fatua crosses were notparticularly promising in this study . The intermediate heritability estimates in crosseswith Stout suggest that some progress can be made when selecting for higher groatprotein percentage if the A. saliva parent is low for this trait . As parental differencesin groat protein percentage diminished, heritable variation in progenies diminished .During the course of this study we also noted that most simple cross progenies fromA. saliva x A . fatua crosses had weak straw and were susceptible to crown rust. Weare attempting to alleviate these deficiencies by backcrossing to the A. saliva parents,912

J . M . REICH AND M . A. BRINKMAN

Table 6 . Weighted average phenotypic correlation coefficients for groat protein percentage, grain yield,and 100-seed weight in F,, F 2 , and F, progenies grown at Madison, Wisconsin in 1979 and 1980 .

Euphytica 33 (1984)

Correlated characteristics Correlation coefficients

F, F 2 F,

1979 1980 1979 1980 1980

Protein vs . grain yield -0.05 0.30* 0.24** 0 .28** -0 .20%Protein vs . 100-seed weight -0.13 0 .10 0.34** 0 .14** 0 .08Grain yield vs . 100-seed weight 0.35** 0 .01 0 .22** 0 .01 0 .08

Euphytica 33 (1984)

GROAT PROTEIN IN AVENA

which have good crown rust resistance and stiff straw. Selection in backcross genera-tions will focus primarily on agronomic traits, with limited emphasis on graot proteinpercentage .

REFERENCES

BRIGGLE, L . W ., Y. POMERANZ & G . S . ROBBINS, 1975 . Protein concentration and amino acid compositionof Avena sterilis L . groats . Crop Sci . 15 : 547-550 .

CAMPBELL, A. R. & K . J . FREY, 1972a . Association between groat protein percentage and certain plantand seed traits in interspecific oat crosses . Euphytica 21 : 352-362 .

CAMPBELL, A. R. & K . J. FREY, 1972b . Inheritance of groat protein in interspecific oat crosses . Can . J .Plant Sci . 52 : 735-742 .

FREY, K . J ., 1976 . Plant breeding in the 1970's : Useful genes from wild plant species . Egypt. J . Genet .Cytol . 5 : 460-482.

FREY, K . J. & J . A. BROWNING, 1971 . Association between genetic factors for crown rust resistance andyield in oats. Crop Sci . 11 : 757-760 .

]WIG, M . M . & H . W. OHM, 1976. Genetic control of protein from Avena sterilis L . Crop Sci . 16 : 749-752 .LYRENE, P . & H . L . SHANDS, 1974 . Groat protein percentage in Avena sativa L . fatuoids and in a fatuoid

x A . sterilis L . cross . Crop Sci . 14 : 765-767 .PETR, F. C . & K . J. FREY, 1966 . Genotypic correlations, dominance, and heritability of quantitative charac-

ters in oats. Crop Sci . 6 : 259-262 .OHM, H . W. & F . L. PATTERSON, 1973. Estimation of combining ability, hybrid vigor, and gene action

for protein in Avena spp . L . Crop Sci . 13 : 55-58 .REICH, J . M ., 1981 . Path analysis, hybrid vigor, and the inheritance of groat protein percentage in Avena

sativa L . x A . fatua L . crosses. M .S . Thesis, Univ . ofWisconsin, Madison, WI.RINES, H . W ., D . D. STUTHMAN, L. W. BRIGGLE, V . L. YOUNGS, H. JEDLINSKI, D . M . SMITH, J . A . WEBSTER& P. G. ROTHMAN, 1980. Collection and evaluation of Avena fatua L. for use in oat improvement. CropSci . 20 : 63-68 .

SPILDE, A. L ., R . S . ALBRECHTSEN & M . D. RuMBAUGH, 1974 . Relationship of protein percent with otherphenotypic characters in interspecific oat crosses . Crop Sci. 14: 767-769 .

SRAON, H . S ., D. L. REEVES & M . D. RUMBAUGH, 1975 . Quantitative gene action for protein content ofoats . Crop Sci . 15 : 668-670 .

STEEL, R . G . D . & J . H . TORRIE, 1980 . Principles and procedures of statistics. 2nd ed . McGraw-Hill, NewYork .

TAKEDA, K . & K . J . FREY, 1979 . Protein yield and its relationship to other traits in backcross populationsfrom an Avena sativa x A . sterilis cross . Crop Sci . 19 : 623-628 .

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