Three winter wheat (Triticum aestivum L.) composite cross populations (CCPs) that had been maintained in repeated parallel populations under organic and conventional conditions from the F5 to the F10 were compared in a two-year replicated field trial under organic conditions. The populations were compared to each other, to a mixture of the parental varieties used to establish the CCPs, and to three winter wheat varieties currently popular in organic farming. Foot and foliar diseases, straw length, ear length, yield parameters, and baking quality parameters were assessed. The overall performance of the CCPs differed clearly from each other due to differences in their parental genetics and not because of their conventional or organic history. The CCPs with high yielding background (YCCPs) also yielded higher than the CCPs with a high baking quality background (QCCPs; in the absence of extreme winter stress). The QCCPs performed equally well in comparison to the reference varieties, which were also of high baking quality. Compared to the parental mixture the CCPs proved to be highly resilient, recovering much better from winter kill in winter 2011/12. Nevertheless, they were out yielded by the references in that year. No such differences were seen in 2013, indicating that the CCPs are comparable with modern cultivars in yielding ability under organic conditions. We conclude that—especially when focusing on traits that are not directly influenced by natural selection (e.g. quality traits)—the choice of parents to establish a CCP is crucial. In the case of the QCCPs the establishment of a reliable high-quality population worked very well and quality traits were successfully maintained over time. However, in the YCCPs lack of winter hardiness in the YCCP parents also became clearly visible under relevant winter conditions.
The challenges of climate change, increasing demand for finite resources, and population growth are calling for a paradigm shift in resource use [
Responding to the continuous restriction of genetic variability in plant breeding, Simmonds [
For the self-pollinating cereals, evolutionary breeding based on the composite cross approach was developed. In evolutionary breeding, heterogeneous, segregating crop populations (composite cross populations, CCPs) [
While genetic variability is expected to decrease in each population over time under the combined effects of drift and selection, overall diversity is supposed to be maintained through the differentiation among populations [
In 2001, three winter wheat CCPs suitable for European growing conditions were created in the UK by the John Innes Centre (JIC, Norwich, UK) in cooperation with the Organic Research Centre (Newbury, UK) [
The variety ‘Bezostaya’, known as high yielding as well as high quality in Russia, was included in both groups Y and Q. A comprehensive analysis of the performance of the individual parental varieties was published by Jones et al. [
After two years of multiplication at two organic and two conventional sites in the south and east of the UK, F
In autumn 2006, enough seeds were available to split the populations one more time. Since then, within each system two Y, two Q, and two YQ populations have been maintained as two parallel populations. This has enabled the comparison of changes in the populations over time within and between systems. Random changes and changes in the populations that occurred due to effects of the environment (e.g. organic vs. conventional growing conditions) can be distinguished. The populations are maintained in separated plots of minimum 100 m
Thus, since the F
In 2011/12 and 2012/13 a field trial was carried out at the University of Kassel comparing the total of twelve winter wheat CCPs in an organically managed field a) to each other and b) to three modern pure line varieties well suited for the local growing conditions. The main questions addressed in the field trial were:
What are the effects of organic versus conventional selection environments on population performance?
What are the effects of genetic background on population performance?
How do the populations perform compared to modern pure line wheat varieties currently popular in organic farming?
To assess morphology and the agronomic performance of the CCPs, straw height, ear length, foot and foliar diseases, yield parameters and baking quality parameters were assessed. The results give an insight into the agronomic performance of CCPs that were shaped over several years in contrasting environments.
The trial was carried out at the Research Station of the University of Kassel in Neu-Eichenberg, located 51
In 2011, enough seed of the F
The trials were carried out in an organic field, the pre-crop in 2011 was canola, in 2012 it was two years of grass-clover. The mean availability of mineral nitrogen (kg N/ha) measured in early spring (BBCH 20) in three layers of soil (0–30, 30–60 and 60–90 cm) was 83.7 kg/ha in total in spring 2012 and 84.0 kg/ha in total in spring 2013. At the flowering stage (BBCH 65) the soil could only be sampled down to a depth of 60 cm, due to very dry soil conditions. Mean availability of mineral nitrogen in total of both depths was 21.6 kg/ha in 2012 and 27.1 kg/ha in 2013. Soil samples were taken and analysed according to the standards of VDLUFA [
The sowing date in 2011 was the 31st of October, in 2012 it was the 10
Growth stages were assessed regularly throughout the season. Straw height and ear length (cm) were measured in 50 randomly chosen stems per plot (BBCH 90) in order to evaluate morphological variation. Straw height was measured from the ground to the start of the ear, ear length was measured from the first full spikelet to the tip without awns.
Foliar diseases caused by fungal pathogens were assessed at BBCH stage 73/75. Non-green leaf area was estimated in % (1–100%). The three leaf levels of flag leaf (F), leaf below flag leaf (F-1) and leaf below F-1 (F-2) were assessed separately at six locations per plot.
To assess foot diseases (
Grain yield on a plot basis was measured in t/ha at 14% moisture content, additionally the thousand kernel weight (TKW) was measured in g at 14% moisture. Ear bearing tillers/m
Protein content (%) was calculated from the nitrogen content of the seeds (N [%]
Hagberg falling number (HFN; sec.; ICC Method no. 107), sedimentation value (Zeleny; ml; ICC Method no. 116), and wet gluten (%; ICC Method no. 106/2) were analysed in the Aberham Laboratories, Großaitingen, DE. HFN was assessed in pooled samples in the first year of the trial and per plot in the second year. Sedimentation value and wet gluten were assessed in pooled samples from the four replications in both years.
Baking volume of test loaves (ml) was assessed using an internal method credited to Aberham Laboratories: test loaves were baked from wholemeal, no ascorbic acid was added but due to very high HFN of some samples the addition of malt flour was necessary to prevent the bread crust form liquefying. Baking volume was assessed per plot in the second trial year only. For a detailed rating system and its translation into a color code of the respective values see Table
Foliar disease severity per plot was calculated as the means per leaf level. Means were weighed 4:3:3 for the flag (F) leaves, the F-1 and F-2 leaves, respectively to account for the greater contribution of the flag leaf to the total dry matter of ripe seeds compared to the lower leaves [
A foot disease severity index (DI) was calculated based on the severity classes as:
where x
The statistical calculations were performed using IBM SPSS Statistics (Version 22). Data were tested for normal distribution of residuals (Shapiro-Wilk-Test and Q-Q-plots) and for homogeneity of variance (Levene test) and transformed if required. When data were normally distributed and variance was homogeneous, a univariate ANOVA with subsequent Tukey-B-Test was calculated where appropriate to find significant differences between group means at p
Where normal distribution was the case but not homogeneity of variance, the Games-Howell post hoc test was used (foliar diseases in both trial years, total incidence of foot diseases in 2011/12, and ear length in both trial years). Linear contrasts were calculated to compare
i) the three groups of populations (YQCCP, QCCP and YCCP),
ii) populations and the reference varieties ‘Achat’, ‘Akteur’, and ‘Capo’,
iii) populations and the mixture, and
iv) CCP
Average temperature during the wheat growing season 2011/12 was 9.7
The distribution pattern of the monthly precipitation, however, showed strong deviations from the long-term average. The average total annual precipitation from 1977 to 1994 was 619 mm, from 2000 to 2013 it was 684 and in 2012 and 2013 it was 792 and 657 mm, respectively. There were very dry periods in November 2011, February and March 2012 and in spring 2013, and some extremely wet months in winter 2011, summer 2012 and May 2013 (Figure
The combination of extremes in winter 2011/12 exposed the plots to a severe winter. After two unusually mild and wet winter months temperatures suddenly dropped at the end of January 2012. Three weeks of black frost with minimum temperatures reaching down to
In mid-February, temperatures increased again and March was warm (average monthly temperature 7.5
Year | No. of frost days with daily minimum temperature below 0 |
No. of frost days with daily maximum temperature below 0 |
Average minimum temperature( |
Average maximum temperature ( |
|
2011 | 20 | 7 |
|
|
|
2012 | 19 | 13 |
|
|
|
2013 | 16 | 9 |
|
|
Disease pressure in both years was low. In both years the dominant disease was
For
In 2012, overall straw length was considerably lower than in 2013 (77.2 cm vs. 90.5 cm, respectively). Overall, the CCPs were significantly shorter than the reference varieties in 2012 but not in 2013 and significantly taller than the mixture of the parental varieties in both years. The QCCPs were always significantly taller than the YCCPs (Table
As expected, within-plot variation of
In 2013, ‘Capo’ was significantly tallest (99 cm), the mix of parental varieties was shortest (65 cm). The two other references were also very short (‘Achat’ and ‘Akteur’ with 86 and 87 cm respectively). While ‘Capo’ was tall or tallest in both years, ‘Achat’ and ‘Akteur’ changed in terms of their ranges in straw length values. While ‘Achat’ and ‘Capo’ were considerably shorter in 2012 than in the year after, absolute height of ‘Akteur’ changed only very little (83 cm in 2012 vs. 87 in 2013) and its change of position in the range of varieties and CCPs is only due to the overall taller plants in 2013.
In the group of CCPs, CY I was the shortest in 2013 (88 cm) as it was in 2012, followed by the three other YCCPs. Again, all YCCPs were shorter than the mean height of plants in the trial, forming a subgroup that was statistically distinguishable from the group of the taller YQCCPs and QCCPs (Figure
Variation in
Year | Comparison group | Straw length [cm] | Ear length [cm] | |||||
1 | 2 | 1 | 2 | p-value | 1 | 2 | p-value | |
2012 | CCPs | References | 77 | 81 | 0.012* | 8.8 | 9.2 | 0.046* |
CCPs | Mixture | 77 | 65 |
|
8.8 | 9.1 | 0.294 | |
YQCCPs | QCCPs | 89 | 80 | 0.589 | 8.8 | 8.6 | 0.432 | |
QCCPs | YCCPs | 80 | 72 |
|
8.6 | 9 | 0.12 | |
YQCCPs | YCCPs | 80 | 72 |
|
8.8 | 9 | 0.415 | |
CCP |
CCP |
77 | 77 | 0.71 | 8.9 | 8.6 | 0.228 | |
2013 | CCPs | References | 93 | 91 | 0.13 | 8.9 | 9.6 | 0.001* |
CCPs | Mixture | 93 | 65 |
|
8.9 | 9.3 | 0.463 | |
YQCCPs | QCCPs | 94 | 95 | 0.269 | 8.9 | 8.9 | 0.977 | |
QCCPs | YCCPs | 95 | 93 |
|
8.9 | 9 | 0.376 | |
YQCCPs | YCCPs | 94 | 93 |
|
8.9 | 9 | 443 | |
CCP |
CCP |
93 | 92 | 0.331 | 9 | 8.8 | 0.133 | |
* Groups differ at p |
The average number of ear-bearing tillers/m
While in the first experimental year no differences between groups could be found apart from a significant difference between CCPs and the mixture, some groups varied considerably in the second year. References formed significantly fewer ears than CCPs. The YCCPs (223 ears/m
In 2012, average yield in the trial was 4.2 t/ha with ‘Akteur’ yielding significantly highest (5.5 t/ha) and the mixture yielding lowest (2.9 t/ha). For all four YCCPs yield was less than the average. In 2013, average yield in the trial was 6.1 t/ha, which was 1.9 t/ha more than in 2012, with CY I (C = conventional) yielding highest (6.7 t/ha) and CYQ II yielding lowest (5.4 t/ha). In this year, the YCCPs yielded above average or just about average while QCCPs and YQCCPs yielded lower or just about average (with the exception of OYQ II (O = organic) which also yielded above average). Differences in yield were, however, not statistically significant in 2013 (Figure
In 2012, the reference varieties yielded significantly higher than the CCPs while in 2013 there was no difference. The mixture yielded significantly less than the CCPs in both years and in 2012 the YCCPs yields were significantly lower than the QCCPs and the YQCCPs. The six CCP
The average TKW was 49.6 g in 2012 (Figure
For the
Year | Comparison group | Ear-bearing tillers/m |
Yield [t/ha] | TWK [g] | |||||||
2012 | 1 | 2 | 1 | 2 | p-value | 1 | 2 | p-value | 1 | 2 | p-value |
CCPs | References | 132 | 130 | 0.852 | 4 | 4.9 |
|
49.4 | 50.2 | 0.148 | |
CCPs | Mixture | 132 | 107 | 0.013* | 4 | 2.9 |
|
49.4 | 46.9 |
|
|
YQCCPs | QCCPs | 132 | 131 | 0.915 | 4.1 | 4.2 | 0.565 | 49.9 | 50.3 | 0.312 | |
QCCPs | YCCPs | 131 | 132 | 0.892 | 4.2 | 3.8 |
|
50.3 | 48 |
|
|
YQCCPs | YCCPs | 132 | 132 | 0.977 | 4.1 | 3.8 | 0.011* | 49.9 | 48 |
|
|
2013 | CCP |
CCP |
129 | 134 | 0.401 | 4 | 4 | 0.532 | 49.8 | 49 | 0.013* |
CCPs | References | 207 | 180 | 0.002* | 6.1 | 6.1 | 0.824 | 48.5 | 49.1 | 0.333 | |
CCPs | Mixture | 207 | 181 | 0.049* | 6.1 | 5.3 | 0.015* | 48.5 | 44.2 |
|
|
YQCCPs | QCCPs | 197 | 202 | 0.607 | 6 | 6 | 0.895 | 48.8 | 48.5 | 0.513 | |
QCCPs | YCCPs | 202 | 223 | 0.026* | 6 | 6.3 | 0.148 | 48.5 | 48 | 0.59 | |
YQCCPs | YCCPs | 197 | 223 | 0.007* | 6 | 6.3 | 0.116 | 48.8 | 48 | 0.245 | |
CCP |
CCP |
207 | 207 | 0.918 | 6.1 | 6.1 | 0.818 | 48.9 | 48 | 0.135 | |
* Groups differ at p |
In general, it could be observed that in both years YCCPs were clearly separate from the other populations and varieties with the YCCPs ranging lowest for all baking quality parameters tested. The QCCPs were in both similar to the reference varieties, which is also consistent for all parameters except protein content in 2013, where QCCPs had a significantly higher protein content than the references. The YQCCPs ranged in both years between the other groups of populations and varieties regarding all values tested and also the finding that CCP
Values for protein content, HFN, baking volume, wet gluten as well as sedimentation value were close to the average of the trial in both years (Table
When comparing groups (Table
For
For
Year | 2012 | 2013 | |||||||
HFN [sec.] | Sedimentation value [ml] | Wet gluten [%]* | Protein content [%] | HFN [sec.]
|
Sedimentation value [ml]* | Wet gluten [%]* | Protein content [%] |
Baking volume [ml] |
|
OYQ I | 275 | 41 | 29.4 | 12.2 |
246 |
30 | 26.7 | 11.2 | 373 |
OYQ II | 293 | 38 | 28.4 | 11.8 |
236 |
31 | 28.4 | 11.7 | 379 |
OQ I | 309 | 51 | 28.5 | 12.5 |
308 |
39 | 27.5 | 11.9 | 428 |
OQ II | 349 | 49 | 28.4 | 12.3 |
325 |
37 | 27.3 | 11.7 | 418 |
OY I | 207 | 29 | 28.5 | 12.2 |
181 |
19 | 25.4 | 11.3 | 350 |
OY II | 206 | 27 | 27.8 | 11.9 |
237 |
20 | 25.8 | 11.1 | 344 |
CYQ I | 274 | 39 | 29 | 12.1 |
284 |
29 | 25.4 | 11.1 | 361 |
CYQ II | 256 | 40 | 29.5 | 12.1 |
291 |
29 | 25.6 | 11.4 | 367 |
CQ I | 307 | 49 | 28.8 | 12.5 |
295 |
37 | 27.6 | 11.7 | 401 |
CQ II | 296 | 47 | 28.6 | 12.5 |
313 |
41 | 28.2 | 11.9 | 413 |
CY I | 204 | 30 | 29 | 12.2 |
203 |
20 | 24.7 | 11 | 359 |
CY II | 219 | 29 | 27.7 | 11.7 |
208 |
22 | 25.4 | 11.1 | 361 |
Achat | 396 | 46 | 26.9 | 11.5 |
370 |
41 | 27.5 | 11.3 | 408 |
Akteur | 424 | 38 | 24.8 | 10.8 |
347 |
34 | 21.6 | 10.1 | 403 |
Capo | 371 | 66 | 31.6 | 12.9 |
392 |
46 | 27.1 | 11.5 | 385 |
Mixture | 302 | 55 | 29.2 | 12.6 |
242 |
30 | 26.7 | 11.3 | 363 |
mean | 292 | 41 | 28.5 | 12.1 | 282 | 32 | 26.3 | 11.3 | 383 |
* Data from pooled samples. | |||||||||
|
Comparison group | Protein content [%] 2012 | Protein content [%] 2013 | Baking volume [ml] 2013 | HFN [sec.] 2013 | |||||||||
1 | 2 | 1 | 2 | p-value | 1 | 2 | p-value | 1 | 1 | p-value | 1 | 2 | p-value |
CCPs | References | 12.2 | 11.7 |
|
11.4 | 11 | 0.118 | 380 | 399 | 0.033* | 260 | 370 |
|
CCPs | Mixture | 12.2 | 12.3 | 0.282 | 11.4 | 11.3 | 0.816 | 380 | 363 | 0.246 | 260 | 243 | 0.574 |
QCCPs | References | 12.4 | 11.7 |
|
11.8 | 11 | 0.016* | 415 | 399 | 0.127 | 310 | 370 | 0.014* |
YQCCPs | QCCPs | 12 | 12.4 |
|
11.3 | 11.8 | 0.144 | 370 | 415 |
|
262 | 310 | 0.042* |
QCCPs | YCCPs | 12.4 | 12 |
|
11.8 | 11.1 | 0.033* | 415 | 354 |
|
310 | 205 |
|
YQCCPs | YCCPs | 12 | 12 | 0.938 | 11.3 | 11.1 | 0.489 | 370 | 354 | 0.104 | 262 | 205 | 0.014* |
CCP |
CCP |
12.1 | 12.2 | 0.812 | 11.5 | 11.4 | 0.629 | 384 | 377 | 0.548 | 256 | 265 | 0.577 |
* Groups differ at p |
Overall, differences due to the parental background of the CCPs and not due to their conventional or organic history were clearly evident in the trials. Compared to the parental mixtures, the CCPs proved to be highly resilient, recovering much better from winter kill in 2012. Nevertheless, they were outyielded by the references in 2012 but not in 2013. In contrast, baking quality of the QCCPs was not different from that of the high baking quality reference varieties.
Disease pressure was low and thus did not play a role for the performance of the CCPs or the references during the two experimental years. Overall, there was neither an influence of the choice of parents nor of the growing system visible. Parents were chosen with the focus on yield and baking quality and not in order to represent different disease resistances, therefore it is unlikely that the CCPs initially differed very much regarding their resistances. Disease pressure in the growing environment where the populations evolved was moderate and did not differ much between the organic and conventional growing area, this meant a strong differentiation of populations was not expected.
Higher disease pressure might have resulted in a different picture as the results of other experiments indicate. Observations of powdery mildew (
Observations in stripe rust (
Since 2011, new races of stripe rust have made a dramatic appearance throughout Europe [
The CCPs as well as the references could not reach their full height potential in the first year due to the extreme weather conditions. The same was reported from regional variety trials, where the average plant height of winter wheat grown without growth regulators in 2012 was reported to be only 87 cm [
The parents were equally short in both years as they were mostly dwarf types. In contrast, the CCPs were much taller indicating that the dwarfing genes have decreased in frequency. They might not have been eliminated completely though, as variation for this trait is still quite large. Nevertheless, the CCPs were within the normal height range; they were shorter than the references in the first experimental year and about the same height in the second year.
Findings of Goldringer
Morphological characteristics of the parental varieties were documented in 2007 [
Measurements in the F
Ear-bearing tillers were at the same low level for all CCPs, the mixture and the references without large variation in summer 2012, which shows that the winter conditions influenced all plots in a similar way resulting in overall low yields. Nevertheless, the resilience of the CCPs and reference varieties was remarkably higher than for most of the parents (Figure
Based on previous year’s results [
A closer look at the pedigree reveals also here a German winter wheat variety—‘Disponent’—as a crossing partner [
The comparably good yield of ‘Achat’, ‘Akteur’, and ‘Capo’ in 2012 is most likely owed to their relatively good winter hardiness and to the fact that good winter hardiness was not one of the main traits in focus when establishing the CCPs. It remains to be seen if the CCPs respond better to freezing after having survived one especially cold winter. As we used the same seed in both years the winter effects did not affect the performance in the second year. Results from experiments investigating the effect of natural selection on the winter survival of barley CCPs indicate that natural selection did increase winter survival although not uniformly over different generations [
In 2013, yield of the YCCPs corresponded with expectations being 0.3 t/ha higher than the QCCPs and YQCCPs. These differences were, however, not statistically significant. Yield of the CCP
A higher TKW was the only parameter that separated the CCP
Characteristics for early vigour were improved after five years in the organically managed CCPs in comparison to the conventionally managed CCPs. The changes towards early vigour in the organic CCPs are thought to be due to the combined effects of selection for higher nitrogen uptake under low-input conditions, and increased competition for light and larger seeds, rather than a direct adaptation to higher weed pressure [
As baking tests are rather costly and time-consuming, various indirect parameters such as sedimentation value, wet gluten, protein content and falling number are often used to predict the baking properties of wheat flour. It has been assumed that protein and wet gluten content strongly correlate with the baking volume determined in the RMT. This is, however, not always the case [
In our study indirect baking quality parameters were analyzed in both years while baking tests could only be conducted in 2013. The results for protein content and HFN in 2013 were in accordance with the outcomes of the baking tests while wet gluten and sedimentation value were less suitable to predict the baking test outcome. The results show a clear differentiation of groups based on the original composition of the populations for all parameters, except wet gluten.
For a wholemeal baking test the average volume of loaves of 383 ml is a good result. Baking with wholemeal flour, lower volumes are the norm and a volume of 400 ml or above is considered very good, 350 to 400 ml is good, below 350 ml is moderate and 330 ml and below is poor (pers. comm. Dr. R. Aberham).
In the test, all CCPs and references except OY II ranged above 350 ml. The strong differences between varieties that can be observed with white flour are less pronounced when testing with wholemeal flour [
While yield is a trait that is subject to natural selection [
The crossing design of the CCPs developed by the John Innes Centre and Elm Farm Research Centre took it into account that quality traits are not subject to natural selection. As opposed to the early composite cross populations of wheat and barley [
Looking at the yield and quality achieved by the mixture of parents a contrast of low yield in both years, but good quality becomes visible. The CCPs out yielded the parental variety mixture in both years. Here the populations seem to have a clear advantage over the mixture. The overall higher diversity and/or natural selection and adaptation over time may be responsible for this. For the quality aspect natural selection played – as mentioned above – a minor role and QCCPs and parents continued to perform similarly after a decade of selection.
The concept of evolutionary breeding can be one of the new, different and efficient strategies urgently required to face the challenges of climate change, population growth and use of finite resources. The overall question if the growing conditions on either organic or conventional fields influence the agronomic performance of the populations, cannot be answered conclusively. The two years were very different, especially regarding the climatic conditions, and many differences were not consistent over both years of the trial.
The parental selection for the CCPs has a much greater influence on their performance than the growing and management conditions to which the populations are subjected. This can be observed with regards to baking quality traits, as well as with morphological parameters, grain yield and yield parameters.
The choice of parents to establish a CCP is crucial, especially when focusing on traits which are not directly influenced by natural selection (for example, quality traits). In the case of the QCCPs the establishment of a reliable high-quality population worked very well and quality traits were successfully maintained over time.
The results clearly indicate that the intercrossing of several pure line varieties does not strongly disconnect their carefully selected traits and much of the originally exhibited characteristics remain (including lack of winter hardiness, for example). The traits present in the parental varieties determine the performance of the CCPs to a considerable degree, even after several years of adaptation to specific growing conditions, so the initial choice of parents suitable for the intended growing conditions should not be underestimated.
As the populations only evolve slowly or not at all in the absence of high selection pressure, which was illustrated by the reactions to foot and foliar diseases, they might be in danger of being outperformed by newly bred wheat varieties after a decade of maintenance and evolution. The frequent integration of well adapted, modern breeding lines into existing CCPs might help to overcome this constraint. Another strategy could be to apply additional human selection such as mass selection for vigour or disease resistance in the context of participatory breeding approaches.
This project was financially supported by the Bundesanstalt für Landwirtschaft und Ernährung (BLE No: 2810OE082) 2011–2014.
Value | Rating | Further differentiation of rating where possible | |
HFN [sec.] |
|
poor | |
180-239 | moderate | ||
240-280 | good | ||
|
poor | ||
Sedimentation value [ml] |
|
poor | |
23-29 | moderate | ||
30-34 | good | good | |
35-40 | good | very good | |
good | Aufmischqualität | ||
Wet gluten [%] |
|
poor | inacceptable |
20-23 | poor | poor | |
24-25 | moderate | poor to moderate | |
26-27 | moderate | moderate | |
28-30 | good | good | |
|
good | very good | |
Protein content [%] |
|
poor | |
10,5-12,5 | moderate | ||
|
good | ||
Baking volume [ml] (wholemeal) |
|
poor | |
330-349 | moderate | ||
350-400 | good | good | |
|
good | very good |