Euphytica 47 : 49-55, 1990 .©1990 Kluwer Academic Publishers . Printed in the Netherlands .

Dough un-mixing time, and the sticky dough problem associated withSr31 wheats

William C. BarnesNSW Agriculture & Fisheries, Agricultural Research Centre, RMB 944, Tamworth, 2340, New South Wales,Australia

Received 7 March 1989 ; accepted in revised form 17 July 1989

Key words: Triticum aestivum, wheat, Sr31, 1BL/1RS translocation, sticky dough problem,dough un-mixing time, mixing tolerance, over-mixing

Summary

The dough handling properties of a number of Sr31 and non-Sr31 wheats were examined in the laboratorytest bake procedure using a National test bake mixer . Dough stickiness was not apparent in any of the wheatsat optimum dough development . The baking quality at optimum dough development of Sr31 wheats wascomparable to non-Sr31 wheats . However, after optimum dough development Sr31 wheats broke down andexhibited dough stickiness more rapidly with continued mixing than non-Sr31 wheats .

A new dough parameter is introduced, dough un-mixing time, and is defined as the time from the point ofoptimum dough development to the point where the dough breaks down as a result of continued mixing toproduce a sticky, non-cohesive mass .

It is shown that Sr31 wheats have shorter dough un-mixing times than non-Sr31 wheats, and that doughun-mixing times for both Sr31 and non-Sr31 wheats are influenced by environmental factors, particularlythose which determine grain protein content .

A selection strategy for breeding Sr31 wheats with commercially acceptable dough properties is indicatedby placing Sr31 in a genetic back-ground of high quality gluten proteins, and selecting for long doughun-mixing times .

Introduction

It is generally accepted that the introduction of ryegenetic material into wheat lowers baking qualitydue to the occurrence of sticky doughs, which causehandling problems during processing (Zeller et al .,1982; Moonen & Zeven, 1984; Payne et al ., 1987) .The phenomenon is shown in Fig . 1, where Sun89D, an Sr31 wheat, exhibits intense dough sticki-ness when mixed in the test bake mixer for threeminutes after optimum dough development,whereas Sunco, a non-Sr31 wheat, when similarlymixed shows no indication of dough stickiness .

The short arm of rye chromosome 1R has beenshown to carry many important genes for resistanceto stem, leaf and stripe rust, powdery mildew, andSeptoria leaf spot (Mettin et al ., 1973 ; CIMMYT,1980; Merker, 1982 ; CIMMYT, 1983) . Resistanceto yellow spot has recently been reported (Rees,1988) .

Tolerance to drought and high grain yield poten-tial over a wide range of environments have alsobeen reported (Rajaram et al ., 1983) .

The stem rust resistant gene Sr31 from Petkusrye, located on 1RS, has been successfully incorpo-

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Fig. 1 . Sunco, a non-Sr31 wheat, and Sun 89D, an Sr31 wheat,mixed for 3 minutes after optimum dough development . Sun89D exhibits the sticky dough phenomenon, whereas Sunco isnot affected .

rated into hexaploid wheats (Zeller, 1973 ; McIn-tosh, 1983) .

Sr31 has been introduced into Australian breadwheats through the 1BL/1RS translocation, but hasbeen precluded due to the association with thesticky dough problem (Martin & Stewart, 1986) .

The cause of the sticky dough problem is notestablished, and research has concentrated on thepossible identification of chemical constituents,such as water soluble proteins and pentosans (Zell-er et al ., 1982 ; Dhaliwal et al ., 1988) .

The objective of this paper was to study theeffects of continued dough mixing on the doughhandling properties of Sr31 and non-Sr31 wheats,in relation to the Sr31 sticky dough problem, whenmixed in a laboratory test bake mixer .

Dough handling properties were examined interms of dough development time, optimum doughdevelopment (Pomeranz, 1987 ; AACC test bakeMethod 10-10A, AACC, 1983), and dough un-mix-ing time .

Dough un-mixing time is a newly introduceddough parameter, and is the time from the point ofoptimum dough development to the point wherethe dough breaks down with continued mixing toproduce a sticky, non-cohesive mass . This is a mea-sure of mixing tolerance, and accordingly a mea-sure of a dough to withstand over-mixing .

A selection strategy is indicated for developingSr31 wheats with commercially acceptable doughproperties, which would be tolerant to over-mix-ing, and not break down and exhibit dough sticki-ness in commercial bakery dough mixers .

Materials and methods

Flour samples

In the preliminary study on the effects of continueddough mixing on the dough handling properties ofSr31 and non-Sr31 wheats the samples used in Fig .2 and Table 1 were obtained from several sources .

Sunco, a strong Prime Hard cultivar, Banks amedium-strong Prime Hard cultivar, and Kite amedium-strong Australian Hard cultivar, were ob-tained as grain samples from Departmental trialsfrom different sites in the 1986 growing season .These were Buhler milled, according to AACCMethod 26-20 (AACC, 1983) .

The two samples of Sun89D, an Sr31 wheat,were obtained as flours from the Bread ResearchInstitute of Australia, North Ryde, New SouthWales, and consisted of a high and a low proteinsample .

The samples of M3344 and M3345, both Sr31wheats, were obtained as flours from the Agricul-tural Research Institute, Wagga Wagga, NewSouth Wales .

In a further study to examine the relationshipbetween dough un-mixing time and the Sr31 stickydough problem, flour samples were obtained undercode from a non-Departmental wheat breedingprogram, courtesy of Dr F . Ellison, Plant BreedingInstitute, Narrabri, New South Wales . These con-sisted of flour samples from two Sr31 wheats and anon-Sr31 control from three different sites, fromthe 1987 wheat growing season .

Flour protein contents were obtained from a Ne-otec 101 (Pacific Scientific, Australia) calibratedagainst Kjeldahl protein (AACC, 1983) .

Test bake dough development curves

These were obtained for the samples shown in Fig .2 and Table 1 . A complete curve is shown in Fig . 2,whereas only the relevant parameters are shown inTable 1 .

To obtain the curve shown in Fig . 2 flours werebaked according to a method similar to AACCMethod 10-IOA (AACC, 1983). Flour weights of

110g (14% m.b .), optimal baking absorption esti-mated from the farinograph (Y = 28 .3 + 0.51 Xwhere Y = baking absorption %, and X = fari-nograph water absorption % ; r = 0 .995), 2% salt,0.1% ammonium chloride, 2 .5% compressedyeast, optimal malt supplementation, and 10p.p.m. potassium bromate were mixed in a Nation-al test bake mixer (National Mfg . Co., Lincoln,Nebraska) operating at 94 r .p.m . for specified mix-ing times .

Doughs were fermented for 2 hours at 28 .3'C,followed by a first punch in a Mono Universal (D .Ayres Jones and Co . Ltd, Swansea, UK), ferment-ed for a further 30 minutes, followed by a secondpunch, followed by a 50 min proof time at 32 .2 ° C .Loaves were baked for 25 min at 232'C .

Doughs were mixed for increasing mixing timesbeginning at 2 min, and proceeding at quarter min-ute intervals . For each mixing time the dough piecewas subsequently baked . This procedure was con-tinued until the dough piece reached a stage ofbreakdown and stickiness such that it could nolonger be handled by the test baker, and thereforecould not be panned and baked. This representedthe un-mixing time end point. Optimum doughdevelopment was determined as being the point onthe dough development curve where loaf volumewas at a maximum .

The results in Table 2 were obtained by a similarprocedure except that the flour formula was re-duced to 110 g of flour, optimal water absorption,and 2% salt . The mixing procedure was continuedas previously until it could no longer be handled bythe test baker, but the dough pieces were notbaked . Dough development time was determinedfrom the farinograph development time using aregression equation, shown in the results section .

Farinographs . These were obtained using the con-stant dough weight method in AACC Method 54-21 (AACC, 1983) .

Results

Figure 2 shows the test bake dough developmentcurve for the strong wheat cultivar, Sunco, a non-

goo

800C)U

7 700

JOu_ 600a0J

500

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1

2 3

4

5 6 7 8 9 10 11 12

DOUGH MIXING TIME, MINS.

Fig. 2. Test bake dough development curve for the strong PrimeHard cultivar, Sunco .

Sr31 wheat . Dough development time is 3 .75 min-utes, and represents the mixing time to achieveoptimum development of the gluten protein ma-trix. At this point maximum gas retention is ob-tained, and therefore loaf volume and loaf qualityare at a maximum . This is represented by point Aon the curve .

Dough un-mixing time is 7 .25 minutes, and is thetime from the point of optimum dough develop-ment to the un-mixing time end point, which isrepresented by point B on the curve . At this pointthe dough is completely broken down into a stickynon-cohesive mass through the continued mixingaction of the mixer . The dough piece cannot behandled by the test baker, and therefore cannot bebaked into a loaf .

Figure 2 therefore shows that non-Sr31 wheatssuch as Sunco will exhibit sticky doughs if they aremixed for a sufficiently long time after the point ofoptimum dough development .

Table 1 shows the data on dough un-mixing timeobtained in a preliminary study on the Sr31 stickydough problem . The loaf volumes shown are thoseobtained at optimum dough development .

The data show that the test loaves baked fromSr31 wheats are comparable to those obtained fromthe Prime Hard cultivars Sunco and Banks . Doughstickiness was not apparent in any of the doughs atoptimum development . Beyond this point doughstickiness became apparent at approximately onehalf of the dough un-mixing time . At the un-mixing

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time end point all doughs were totally un-cohesive,and could not be handled by the test baker .

The data indicate that Sr31 wheats have consid-erably shorter dough un-mixing times than non-Sr31 wheats . The theoretical reasons for this will bediscussed later . Consequently, Sr31 wheats lackmixing tolerance, and as a result break down morerapidly in the dough mixer to produce stickydoughs. On this basis, the short un-mixing timesassociated with Sr31 wheats would appear to berelated to the Sr31 sticky dough problem .

An important result is that obtained for M3345 .Whilst the un-mixing time for this Sr31 wheat isshort, it is comparable to the non-Sr31 cultivarKite, which has been in commercial production formany years without any criticisms of dough sticki-ness. This would indicate that it may be possible toproduce Sr31 wheats with dough mixing tolerancescomparable to commercially acceptable cultivars .

The results presented in Table 2 are the results ofa study consisting of two different Sr31 wheats anda non-Sr31 wheat control, from three differentsites . These samples were received under codefrom a non-Departmental wheat breeding pro-gram. In this study the samples were not baked, butotherwise mixed and doughs handled according tothe test bake dough development curve procedureas shown in Fig. 2. Because the doughs were notbaked, dough development times were determinedfrom the correlation (r = 0 .975) between test bakedough development times (Y) and farinographdough development times (X) using the regressionequation Y = 0.774 + 0.36X. This regression isused routinely in the Tamworth test bake proce-dure, where all doughs are mixed to optimum de-

Table 1 . Data on dough un-mixing times obtained in a preliminary study on Sr31 wheats from various locations

velopment, similar to the AACC test bake Method10-10A (AACC, 1983) .These results were subsequently confirmed by

the wheat breeder as being correct for both wheatvariety and site . Sr31A had the shortest un-mixingtime over all three replicates, and was significantlydifferent from the non-Sr31 wheat . Sr31B had anun-mixing time longer than Sr31A, but shorter thanthe non-Sr31 wheat, and whilst not significantlydifferent from either, it does indicate the possibilityof selecting for longer un-mixing times in Sr31wheats .

Dough un-mixing time is a measure of mixingtolerance, and a certain minimum dough unmixingtime is required for a dough not to exhibit doughstickiness in a commercial bakery . Under commer-cial bakery conditions, particularly where highspeed mechanical dough developers are used, it isnot always possible to precisely control dough mix-ing in terms of dough development, and should thedough be overmixed beyond optimum develop-ment, a critical minimum dough un-mixing time isnecessary to ensure the dough does not break downand become sticky .

A very pronounced site-environmental effect ondough un-mixing time is apparent . The site meanswere site 1- 6.9 mins, site 2 - 2.3 mins and site 3 -12 .3 mins, with a least significant difference (5%)of 7.2 mins. This effect indicates that a control non-Sr31 wheat must be used at each site to assessenvironmental variation . The data also reveal thatnon-Sr31 wheats under certain environmental con-ditions can have relatively short unmixing times asshown by the non-Sr31 wheat at site 2, and this may

Wheat Flour protein %(14% m.b .)

Loaf volume c .c . Development time,mins

Un-mixing time,mins

Sunco, non Sr31 12 .2 815 3 .75 7.25Banks, non Sr31 11 .7 815 2 .25 4.75Kite, non Sr31 12 .3 750 2.75 3.50Sun 89D, Sr31 11 .8 730 2.75 2.25Sun 89D, Sr31 15 .5 850 3.25 2.00M3344, Sr31 11 .5 810 2.75 2.25M3345, Sr31 11 .3 805 3 .25 3.75

account for the occasional occurrence of stickydoughs in non-Sr31 wheats .

Discussion

A test bake dough development curve as shown inFig . 2 characterises a particular flour, and dependson a number of factors such as the dough strengthof the cultivar, protein content and starch damage .This curve is a better representation of the mixingcharacteristics of a flour than that obtained fromthe farinograph or mixograph (AACC, 1983) be-cause it is obtained directly from the test bakemixer, in this case, a National laboratory mixer .Dough stickiness as a result of continued mixingbreakdown could not be obtained in the farino-graph or mixograph, presumably due to the weakermixing action, as compared to the much strongermixing action of the National mixer . It is intendedin the near future to attach a transducer and record-er to the National mixer to obtain an objectivegraphical representation of the dough develop-ment curve. It is not necessary to actually bake thedough pieces, because the height of the graphwould indicate dough development, and thechange in width of the recorded trace from compar-atively wide to narrow would provide an objectivemeasurement of the un-mixing time end point .

The graph in Fig . 2 indicates that when interpre-tating the mixing properties of a particular flour it

Table 2 . Data on dough un-mixing times of Sr31 and non-Sr31 wheats at three locations

should be done in terms of a dough developmentcurve performed in the dough mixer from whichthe bread is to be baked . This would apply equallyto both laboratory mixer and commercial bakerymixers. Mixers such as farinographs and mixo-graphs are only simulations of the real situation .The handling properties of a dough should al-

ways be assessed in relation to the stage of doughdevelopment of the particular dough . Comparisonsbetween doughs of different cultivars should al-ways be done at comparable points on the doughdevelopment curve . To compare the handlingproperties of different doughs at the same mixingtime from zero is erroneous, as one dough mayhave a short development time and be over mixed,whereas the other dough may have a longer devel-opment time and be at optimum development, andtherefore have good dough handling properties .Figure 1 shows two doughs at three minutes mixingafter the point of optimum development . The Sr31dough is completely un-mixed and exhibits intensedough stickiness, whereas the non-Sr31 dough isnot un-mixed and does not exhibit stickiness .

The results shown in Tables 1 and 2 indicate thatSr31 wheats tend to have short dough un-mixingtimes, whereas non-Sr31 wheats have longer doughun-mixing times . Because of the short dough un-mixing times, Sr31 wheats lack mixing tolerance,and break down more rapidly in the mixer to pro-duce sticky doughs . However, the results indicate itmay be possible to select for longer un-mixing

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Wheat Site Flour protein %(14% m.b .)

Development timemins

Un-mixing timemins

Mean un-mixing timemins

Sr31A 1 13 .5 3.50 1 .25 1 .92Sr31A 2 12 .00 3.00 0.50Sr31A 3 7.8 2.00 4.00

Sr31B 1 13 .9 7.00 6.00 6 .58Sr31B 2 10 .4 2 .25 1 .75Sr31B 3 7 .6 2.00 12.00

Non-Sr31 1 13 .0 4.00 13 .50 13 .00Non-Sr31 2 12 .2 3 .75 4.75Non-Sr31 3 8 .3 2 .25 20.75

Least significant difference (5%) of means 7 .2

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times in Sr31 wheats to obtain un-mixing timescomparable with non-Sr31 wheats .

The strong environmental effect demonstratedin Table 2 may perhaps be due to the increase in thegliadin glutenin ratio as grain protein increases(Doekes & Wennekes, 1982) . This would tend todilute the proportion of strong inter-molecular dis-ulphide linkages in the developed dough proteinmatrix, and increase the proportion of hydrogenbonds, electrostatic and ionic bonds which are con-siderably weaker inter-molecular linkages (Pom-eranz, 1987). Consequently, dough un-mixingtimes would tend to be shorter at higher proteinlevels. This may also explain the frequent observa-tion that dough stickiness is more apparent at high-er protein levels in Sr31 wheats . It may also accountfor the occasional occurrence of sticky doughs innon-Sr31 wheats . Environmental factors duringgrain ripening may also influence dough un-mixingtimes through the effects of moisture availabilityand ambient temperatures on the gliadin gluteninratio, and this is currently under investigation .The dough un-mixing phenomenon discussed

here is not unlike that discussed by Tipples & Kil-born, 1975, who found a marked deterioration indough and bread properties when a high speeddeveloped dough was over-mixed at slow speedbeyond a certain stage of breakdown and sticki-ness .

To account for the short dough un-mixing timesof Sr31 wheats it is hypothesised that the replace-ment of the short arm of the 1B chromosome by theshort arm of the 1R chromosome results in theincorporation of relatively large levels of secalins(Lawrence & Shepherd, 1981) in the flour. Thisqualitative observation is supported by acid(pH 3 .1) electrophoresis in polyacrylamide gels of70% aqueous ethanol soluble grain proteins (Bush-uk & Zillman, 1978) . The levels of low molecularweight glutenin sub-units, plus y- and w-gliadins arereduced by one third due to the absence of the shortarm of chromosome 1B (Payne, 1987) . According-ly, the addition of rye secalins, and the loss ofwheat prolamins could be expected to have an ad-verse effect on dough properties .

Whilst the secalins are probably bound into theaggregated gluten protein matrix during dough de-

velopment, they would not be expected to interactwith the glutenin proteins to the same degree asgliadin proteins, because gluten is not easily madefrom rye flours (Shewry & Miflin, 1985) . Theweaker inter-molecular aggregation properties ofthe secalins would result in a more rapid break-down with mixing beyond optimum dough devel-opment. This may account for the absence ofdough stickiness up to and at optimum develop-ment, and the manifestation of dough stickinessafter optimum development .

On the basis of the results presented here doughun-mixing time would appear to be an importantparameter influencing the Sr31 sticky dough prob-lem. It provides an objective measurable doughparameter to enable the evaluation of this prob-lem . Reports of satisfactory dough handling prop-erties in 1BL/IRS translocations such as Pitomaand several new cross-bred wheats (Jost, 1988) maypossibly be accounted for on this basis .

A selection strategy for breeding Sr31 wheatswith satisfactory dough handling properties wouldbe to screen for the high quality high molecularweight glutenin subunits on the long arms of wheatchromosomes 1A, 1B, 1D (Day et al ., 1986), to-gether with good quality low molecular weight glu-tenin sub-units, and y- and w-gliadins on the shortarms of 1A and 1D plus the a- and (3-gliadins on theshort arms of 6A, 6B and 6D (Pogna et al ., 1982 ;Campbell et al., 1987) . Because of the extensiveallelic variation at these major prolamin gene locithe potential genetic variation for dough qualityrelated proteins in a genotype is considerable(Payne, 1987) . Further selection would be based ondough un-mixing time to ensure the un-mixing timeis sufficiently long such that dough stickiness wouldnot be apparent under commercial bakery condi-tions. Such a wheat would have good mixing toler-ance, and acceptable commercial baking perform-ance .

Acknowledgements

The assistance of Ms S . Balfe, test baker, is grate-fully acknowledged .

References

AACC, 1983 . Experimental milling-Buhler method, Method26-20 ; Kjeldahl protein, Method 46-12 ; test bake, Method10-10A; farinograph, constant dough weight procedure,Method 54-21 ; mixograph, Method 54-40 . In : ApprovedMethods, American Association of Cereal Chemists Inc ., St .Paul, Minnesota, USA, 8th edn .

Bushuk, W. & R.R. Zillman, 1978 . Wheat cultivar identifica-tion by gliadin electrophoresis . I . Apparatus, method, andnomenclature . Can . J . Plant Sci . 58 : 505-515 .

Campbell, W.P ., C.W. Wrigley, P .J. Cressey & C .R. Slack,1987 . Statistical correlations between quality attributes andgrain protein composition for 71 hexaploid wheats used asbreeding parents . Cereal Chem . 64 : 293-299.

CIMMYT, 1980 . Probing the gene pools - spring x wintercrosses in bread wheat . CIMMYT Today No . 12 . Internation-al Maize and Wheat Improvement Centre, Mexico City, Mex-ico, 1980 .

CIMMYT, 1983 . Research highlights . International Maize andWheat Improvement Centre, Mexico City, Mexico, 1984 .

Day, P.R ., J . Bingham, P.I. Payne & R.D. Thompson, 1985 .The way ahead : wheat breeding for quality improvement . In :J.M.V. Blanshard, P .J . Frazier & T. Galliard (Eds), Chem-istry and Physics of Baking, pp 251-261 . Special publicationNo. 56, The Royal Society of Chemistry, London, 1986 .

Dhaliwal, A.S ., D .J . Mares, D .R. Marshall & J.H. Skerrit,1988 . Protein composition and pentosan content in relation todough stickiness of 1B/1R translocation wheats . CerealChem. 64 : 72-76 .

Doekes, G .J. & L.M.J. Wennekes, 1982 . Effect of nitrogenfertilization on quantity and composition of wheat flour pro-tein . Cereal Chem . 59 : 276-278 .

Jost, M ., 1988 . Sticky dough related to 1B/1R translocation .Annual Wheat Newsletter, V 34 p 170 . University Communi-cations, Colorado State University, Colorado, USA .

Lawrence, G .J . & K.W. Shepherd, 1981 . Chromosomal locationof genes controlling seed proteins in species related to wheat .Theor. Appl . Genet . 59: 25-31 .

Martin, D .J . & B .G . Stewart, 1986 . Dough mixing properties ofa wheat-rye derived cultivar . Euphytica 35: 225-232 .

McIntosh, R.A ., 1983 . A catalogue of gene symbols for wheat(1983 edition) . In : Proc 6th Int . Wheat Genetic Symp ., pp1197-1254. Kyoto, Japan .

Merker, A ., 1982 . 'Veery' a CIMMYT spring wheat with the

55

1B/1R chromosome translocation . Cereal Res . Comm. 10:105-106 .

Mettin, D ., W.D . Bluthner & G . Schlegel, 1973 . Additionalevidence on spontaneous 1B/1R wheat-rye substitutions andtranslocations . In : Proc 4th Int . Wheat Genet . Symp ., pp179-184 . Columbia, MO, USA .

Moonen, J.H.E. & A.C. Zeven, 1984 . SDS-PAGE of the highmolecular weight subunits of wheat glutenin and the charac-terisation of 1R(1B) substitution and 1BL/1RS translocationlines . Euphytica 33 : 3-8 .

Payne, P .I ., M.A. Nightingale, A .F . Krattiger & L .M. Holt,1987 . The relationship between HMW glutenin subunit com-position and the breadmaking quality of British-grown wheatvarieties . J . Sci . Food Agric . 40 : 51-65 .

Payne, P .I ., 1987 . Genetics of wheat storage proteins and theeffect of allelic variation on bread-making quality . Ann . Rev .Plant Physiol . 38 : 141-153 .

Pogna, N.E ., G . Boggini, M . Corbellini, M . Cattaneo & A . DalBelin Peruffo, 1982 . Association between gliadin electropho-retic bands and quality in common wheat . Can . J . Plant Sci .62: 913-918 .

Pomeranz, Y. ,1987 . Chemical bonds in dough . In : Y. Pomeranz(Auth), Modern Cereal Science and Technology, pp 173-181 .VCH Publishers Inc ., New York, USA .

Rajaram, S ., C .E . Mann, G . Ortiz-Ferrara & A . Mujeeb-Kazi,1983 . Adaptation, stability and high yield potential of certain1B/1R CIMMYT wheats . In : Proc 6th Int . Wheat Genet .Symp., pp 613-621 . Kyoto, Japan .

Rees, R.G ., 1988 . Resistance to yellow spot (Pyrenophora tri-tici-repentis) in wheat . In : 4th Int . Cong . P1 . Path ., Mel-bourne, Australia, abs 823 .

Shewry, P .R. & B .J . Miflin, 1985 . Seed storage proteins ofeconomically important cereals . In : Y. Pomeranz (Ed), Ad-vances in Cereal Science and Technology, Vol VII, pp 1-83 .American Association of Cereal Chemists Inc ., St . Paul,Minnesota, USA .

Tipples, K .H. & R.H. Kilborn,1975 . `Unmixing'-the disorien-tation of developed bread doughs by slow speed mixing .Cereal Chem . 52: 248-262 .

Zeller, F .J ., 1973 . 1B/1R wheat-rye chromosome substitutionsand translocations . In : Proc 4th Int. Wheat Genet . Symp . pp209-221 . Columbia, MO, USA.

Zeller, F .J ., G. Gunzel, G . Fischbeck, P. Gerstenkorn & D .Weipert, 1982 . Veranderungen der Backeigenschaften desWeizens durch die Weizen - Roggen - Chromosomen Trans-lokation 1B/1R . Getreide Mehl and Brot 36 : 141-143 .