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Article history: The effect of cooking and dehulling on nutrients and anti-nutritional factors of several varieties of lentils
Received 29 May 2008 (Lens culinaris) was investigated. Significant (p < 0.05) variations existed among the lentil varieties with
Received in revised form respect to their crude protein, starch, ash, soluble dietary fiber (SDF), insoluble dietary fiber (IDF), total
6 October 2008
dietary fiber (TDF), resistant starch (RS), trypsin inhibitor activity (TIA), minerals, phytic acid, tannins,
Accepted 10 October 2008
sucrose and oligosaccharides (raffinose, stachyose and verbascose) content. Cooking lentils in boiling
water significantly increased protein, starch, IDF, TDF, resistant starch, Ca, Cu and Mn content, whereas
Keywords:
reduced ash, Fe, K, Mg, P, Zn, TIA, phytic acid, tannins, sucrose and oligosaccharides were observed.
Lentil
Dietary fiber Dehulling (removal of seed coat) resulted in a significant increase in protein, starch, resistant starch, K, P,
Cooking phytic acid, stachyose and verbascose content, however, a significant decrease in SDF, IDF, TDF, Ca, Cu, Fe,
Dehulling Mg, Mn and tannin content was observed.
Nutrients Crown Copyright Ó 2008 Published by Elsevier Ltd on behalf of Swiss Society of Food Science and
Technology. All rights reserved.
0023-6438/$34.00 Crown Copyright Ó 2008 Published by Elsevier Ltd on behalf of Swiss Society of Food Science and Technology. All rights reserved.
doi:10.1016/j.lwt.2008.10.007
N. Wang et al. / LWT - Food Science and Technology 42 (2009) 842–848 843
cooking and dehulling affect the nutritional and the anti-nutri- a certain amount of water was added to the container to bring the
tional components of several varieties of lentils. moisture content of the sample to 12.5 g/100 g wb. The container
was sealed and kept at 4 C for 3 days with occasional shaking to
2. Materials and methods allow the moisture to equilibrate before the sample was dehulled.
The tempered sample was processed in the mill operated at 1100 r/min
2.1. Materials for 38 s. After dehulling, the processed lentils were separated into
lentil seeds (hulled and dehulled), hulls, powder and broken seeds
Samples were selected from the Canadian Grain Commission using a Carter dockage tester (Simon-Day Ltd, Winnipeg, Canada).
(CGC)’s 2006 harvest survey of the commercially grown crop of The lentil seeds were then separated by hand into their respective
lentils for this study. All the selected samples used in this study hulled and dehulled seeds. Dehulled and raw seeds were ground
were registered varieties and from the same growing region in into flour as described previously and stored at 4 C until further
Saskatchewan, Canada. The eight registered lentil varieties chosen analysis.
were Laird, CDC Sovereign, CDC Richlea, CDC Vantage, Eston,
Milestone, CDC Robin and CDC Blaze which represent the most 2.2.2. Chemical analysis
common varieties grown in Western Canada. Each varietal sample 2.2.2.1. Chemical analysis of nutritional constituents. Nitrogen (N)
was composed of sub-samples (n ¼ 4–8) of a specific registered was determined by the Dumas combustion method using a Leco FP-
variety grown in the same growing region in Saskatchewan. 528 combustion nitrogen analyzer (Leco Corp. St. Joseph, MI, USA)
Samples were cleaned to remove foreign material and damaged calibrated with EDTA (AOAC, 2000). Crude protein content was
seeds prior to processing of lentils (cooking and dehulling). calculated using N 6.25. Moisture and ash content were deter-
Benzoyl-DL-arginine-p-nitroanilide hydrochloride (BAPNA), mined gravimetrically in accordance with AACC methods 44-17 and
trypsin, 5-sulphosalicylic acid, ferric chloride hexahydrate, celite 08-16, respectively (AACC, 2000). Starch was determined colori-
(acid washed), sodium chloride and sodium sulphate were metrically as described by the method AACC 76-13 (AACC, 2000).
obtained from Sigma–Aldrich (Oakville, ON, Canada). Glucose Resistant starch was measured by the AACC method 32-40 (AACC,
hexokinase and amyloglucosidase were purchased from Roche 2000). Soluble, insoluble and total dietary fiber contents were
Diagnostics (Laval, QC, Canada). Resistant starch kit, total dietary determined by sequential enzymatic digestion according to AACC
fiber kit and protease were purchased from Megazyme (Co. Wick- method 35-05 (AACC, 2000). Minerals were determined by atomic
low, Ireland). Sodium Hydroxide (50% w/w) was obtained from absorption spectrophotometry (Gawalko, Nowicki, Babb, & Tka-
Fisher Scientific (Ottawa, ON, Canada). chuk, 1997).
The filtrate (25 ml) was injected on a PA1 analytical anion exchange cooked lentils may be attributed to the loss of soluble solids during
column (4 250 mm) (Dionex Corp., Sunnyvale, CA) connected to cooking, which would increase the concentration of protein and
a CarbopacÔ PA1 guard column (4 50 mm) (Dionex Corp., Sun- starch in cooked seeds. The decrease in ash content might result
nyvale, CA). The mobile phase used was 0.15 mol/L sodium from diffusion of certain minerals into the cooking water. Hay-
hydroxide at the flow rate of 1.0 ml/min. towitz and Matthews (1983) reported that cooking in boiling water
caused great losses in minerals for cooked pulses. Similar results
2.2.3. Statistical analysis have been reported by Wang et al. (2008) for field peas. Dehulling
Two-way analysis of variance (ANOVA) models with variety and resulted in a significant (p < 0.05) increase in protein and starch
treatment (cooking and dehulling) as the model factors were used content for all the varieties (Table 2), confirming results reported by
to analyze the data with the Statistical Analysis System (v.9.1.3, SAS Wang (2008). Since seed coats reportedly contain little protein and
Institute, Cary, NC). A variety-by-treatment interaction was starch, it is opined that dehulled seeds would proportionately
included in the models. A type I error rate (alpha level) of 0.05 was contain more protein and starch. Dehulling had little effect on ash
used for all tests. The Duncan’s multiple range test was used to content in the eight lentil varieties (Table 2).
separate means and significance was accepted at p < 0.05 (Cochran Analysis of variance showed that both variety and cooking had
& Cox, 1992). a significant effect on soluble dietary fiber (SDF), insoluble dietary
fiber (IDF), total dietary fiber (TDF) and resistant starch (RS) content
3. Results and discussion (Table 1). Variety-by-cooking had a significant effect on IDF
(p < 0.05), TDF (p < 0.05) and RS (p < 0.001) but had no effect on
3.1. Effect of cooking and dehulling on proximate composition SDF (Table 1). Table 3 shows values of SDF, IDF and TDF as affected
by cooking and dehulling. SDF content decreased significantly
Analysis of variance showed that variety had a significant (p < 0.05) in the cooked lentils compared to the raw lentils. The
(p < 0.001) effect on protein, starch and ash content (Table 1). results are in fair agreement to those reported by Vidal-Valverde
Similar results have been reported by Wang and Daun (2006). and Frias (1991) who suggested that a softening of soluble fibers
Cooking and dehulling also exhibited a significant (p < 0.001) effect occurred with the cooking process, reducing its content. These
on protein, starch and ash content (Table 1). The interactive effect of results are also comparable to that reported by Wang et al. (2008).
variety-by-cooking on ash content was significant (p < 0.001) However, Kutos, Golob, Kac, and Plestenjak (2003) found that SDF
whereas there was no effect of variety-by-cooking on protein and for beans was increased on cooking. The lowest content of SDF for
starch content (Table 1). Crude protein content (N 6.25) in lentils cooked lentils was determined in the varieties Richlea and
varied from 251.5 to 292.5 g/kg dry matter with a mean of 269.4 g/ Milestone (12.7 g/kg dry matter) and the highest SDF content was
kg dry matter (Table 2). Starch content ranged from 460.1 to in the variety Eston (16.5 g/kg dry matter) (Table 3). Cooking
497.3 g/kg dry matter. Results found in our study were within the significantly (p < 0.05) increased IDF content for all lentil varieties
range reported (Bhatty, 1984; Reddy et al., 1984; Wang & Daun, (133.9–161.1 g/kg dry matter) as compared to the raw samples. This
2006). Cooking resulted in a significant (p < 0.05) increase in crude increase may be due to protein–fiber complexes formed after
protein and starch content but ash content was reduced signifi- possible chemical modification induced by the cooking of dry seeds
cantly (p < 0.05) (Table 2). The increases in protein and starch in (Bressani, 1993). Heat treatment (cooking) may also modify the
Table 1
Analysis of variance of variety and treatment (cooking and dehulling) on composition of lentils.
Variety (V) Cooking (C) V-by-C Variety (V) Dehulling (D) V-by-D
Composition
Protein 1177*** 458.3*** – 1256*** 1346*** 11.4***
Starch 603.6*** 4086*** – 691.3*** 20513*** –
Ash 6.70*** 414*** 1.69*** 4.18*** 2.01*** –
Soluble dietary fiber (SDF) 11.1* 61.9*** – 7.3* 263*** –
Insoluble dietary fiber (IDF) 144.8*** 4357*** 56.2* 34.8* 24647** –
Total dietary fiber (TDF) 122.5*** 33372*** 51.9* 34.7* 30037*** –
Resistant starch (RS) 181.4*** 2188*** 145.5*** 576.0*** 45.6*** –
Minerals
Ca 182.8*** 292.2*** – 86.3*** 13534*** 19.7**
Cu 0.05*** 0.11*** – 0.03*** 0.22*** –
Fe 2.22*** 8.32*** 0.34* 1.25*** 14.2*** 0.94***
K 6248* 855432*** – 4798* 24475** –
Mg 111.0** 960.3*** – 119.3*** 3767*** 33.7*
Mn 0.12*** 0.04* – 0.10*** 0.18** –
P 4150** 10841** – 4215*** 6429** –
Zn 0.72*** 0.54** – 0.66*** – –
Sugars
Sucrose 39.2*** 1579*** 4.94*** 50.5*** 9.79*** 1.02**
Raffinose 0.72*** 51.0*** 0.52*** 1.05*** 0.85*** 0.17*
Stachyose 12.1*** 528.1*** 6.39*** 14.3*** 45.6*** 2.29***
Verbascose 2.48*** 3.25*** 0.24* 2.39*** 26.1*** 0.18*
TIAa 0.18*** 24.5*** 0.06*** 0.15*** 23.9*** 0.08***
Phytic acid 4.11*** 3.45*** 0.11** 4.33*** 1.62*** –
Tannins 1.28*** 32.2*** 1.19*** 1.06*** 173.4*** 1.06***
***, **, * ¼ Significant at p < 0.001, p < 0.01 and p < 0.05, respectively. –; Blank spaces indicate no significance.
a
Trypsin inhibitor activity.
N. Wang et al. / LWT - Food Science and Technology 42 (2009) 842–848 845
Table 2
Effect of cooking and dehulling on protein, starch and ash content of lentils.
Variety Protein (N 6.25) (g/kg dry matter) Starch (g/kg dry matter) Ash (g/kg dry matter)
structure of both cell wall and storage polysaccharides of pulses were significant (p < 0.05) in the dehulled lentils between varieties
possibly by affecting the intactness of tissue histology and dis- studied (Table 3). The percentage of SDF to TDF in the dehulled
rupting the protein–carbohydrate integration, thus reducing the samples ranged from 11.4 to 16.5%, which was significantly
solubility of dietary fiber (Siljestrom et al., 1986). A significant (p < 0.05) higher than in the raw (10.8–13.9%). The ratios of IDF to
(p < 0.05) increase in IDF content in cooked lentils as compared to TDF in the dehulled samples were from 83.5 to 86.9%, which were
their raw forms (Table 3) may be beneficial as the cellulose content significantly (p < 0.05) lower than in the raw (86.9–89.4%). Resis-
of foods in insoluble fiber has been shown to be closely related to tance starch content in dehulled lentils was significantly (p < 0.05)
their glycemic responses (Selvendran, 1984). Hence, it is possible higher than that in the raw lentils (Table 3).
that cooked lentils may still be effective in reducing the glycemic
response despite the reduction in SDF. Cooking increased TDF in 3.2. Effect of cooking and dehulling on mineral content
lentils significantly (p < 0.05) (Table 3). The TDF contents in the raw
lentils were in the range of 131.1–147.0 g/kg dry matter, while TDF A significant varietal effect on calcium (Ca), copper (Cu), iron
varied from 149.1 to 168.7 g/kg dry matter for cooked lentils. The (Fe), potassium (K), magnesium (Mg), manganese (Mn), phos-
IDF fraction in raw lentils formed the majority of TDF, ranging from phorus (P) and zinc (Zn) was found (Table 1), confirming the
86.1% to 89.4%, while the fraction of SDF ranged from 10.6% to 13.9%. findings reported by Wang and Daun (2006). Results showed that K
These results are similar to that reported by Su and Chang (1995) was the most abundant element in raw lentils ranging from 843.5
who reported that the IDF fraction in raw dry beans was 72–90% of to 943.0 mg/100 g (Table 4). P in raw lentils was found to range
the TDF. Wang et al. (2008) observed that IDF was from 88.6% to from 308.0 to 407.0 mg/100 g. Mg varied from 99.3 to 109.0,
90.2% of the TDF for field peas and SDF 9.5% to 11.1% of the TDF. Ca from 57.5 to 76.2, Fe from 6.25 to 9.18 and Zn from 2.63 to
Cooking significantly (p < 0.05) increased resistant starch (RS) 3.79 mg/100 g. Cu ranged from 0.79 to 1.13 and Mn from 1.00 to
content from 15.6–43.2 g/kg dry matter in the raw lentils to 37.3– 1.50 mg/100 g. The mineral contents were in the range reported by
50.9 g/kg dry matter in the cooked lentils (Table 3). The highest Wang and Daun (2006) for lentils and were also comparable with
value in cooked lentils was determined in the variety Richlea those reported for other pulses (Jagadi, Rundgren, & Ogle, 1987;
(50.9 g/kg dry matter). The increase in RS content after cooking is Kadam et al., 1989). Analysis of variance showed that cooking
due to retrogradation of starch after gelatinization (Aman & affected all minerals analysed (Table 1). The interactive effect of
Westerlund, 1996; Raben et al., 1994; Costa et al., 2006). variety-by-cooking on Fe was significant (p < 0.05) but no signifi-
Analysis of variance showed that both variety and dehulling had cant effect was found on other minerals. Cooking lentils in boiling
a significant effect on fiber SDF, IDF, TDF and RS content (Table 1). water resulted in a significant (p < 0.05) increase in Ca, Cu and Mn.
There was no significant effect of variety-by-dehulling on SDF, IDF, However, cooking resulted in significant (p < 0.05) losses of Fe, K,
TDF and RS. SDF contents in the dehulled seeds for the eight lentil Mg, P and Zn (Table 4). Similar results have been reported by
varieties were significantly (p < 0.05) lower than in the raw lentils Haytowitz and Matthews (1983). Dehulling had a significant
(Table 3). Dehulled variety Sovereign had the highest SDF content (p < 0.05) effect on minerals except for Zn (Table 1). Interaction of
(13.4 g/kg dry matter) whereas variety Milestone had the lowest variety-by-dehulling significantly affected Ca (p < 0.001), Fe
SDF (9.5 g/kg dry matter). Dehulling reduced IDF and TDF signifi- (p < 0.001) and Mg (p < 0.05) levels (Table 1). Dehulling signifi-
cantly (p < 0.05) in all eight lentil varieties (Table 3). The decreases cantly (p < 0.05) decreased Ca, Cu, Fe, Mg and Mn (Table 4). Similar
observed in TDF content due to dehulling resulted mainly from results were reported by Singh, Rao, Seetha, and Jambunathan
a decrease in IDF content. Differences in SDF, IDF and TDF contents (1989) for pigeonpea and Wang et al. (2008) for field peas. K and P
Table 3
Effect of cooking and dehulling on soluble, insoluble, total dietary fiber and resistant starch content of lentils.
Variety Soluble fiber (g/kg dry matter) Insoluble fiber (g/kg dry matter) Total dietary fiber (g/kg dry matter) Resistant starch (g/kg dry matter)
Raw Cooked Dehulled Raw Cooked Dehulled Raw Cooked Dehulled Raw Cooked Dehulled
Laird 18.0ad 14.8b 11.6c 123.2b 152.0a 69.2c 141.1b 166.8a 80.8c 33.3c 43.6a 37.6b
Sovereign 17.0a 13.8b 13.4c 114.1b 143.1a 67.8c 131.1b 156.8a 81.2c 15.6c 41.4a 17.9b
Richlea 15.2a 12.7b 11.2c 127.7b 156.0a 63.6c 142.9b 168.7a 74.7c 43.2c 50.9a 47.1b
Vantage 16.1a 13.1b 10.0c 123.5b 144.8a 66.6c 139.6b 158.1a 76.6c 28.1c 41.6a 34.1b
Eston 20.0a 16.5b 11.9c 123.7b 143.5a 69.7c 143.7b 160.5a 81.6c 16.4c 39.7a 21.9b
Milestone 15.6a 12.7b 9.5c 128.3b 161.1a 73.5c 143.9b 173.7a 83.1c 16.0c 40.9a 20.3b
Robin 19.4a 16.0b 12.7c 127.7b 140.9a 67.7c 147.0b 157.0a 80.4c 17.7c 40.8a 21.0b
Blaze 16.7a 15.1b 11.4c 120.4b 133.9a 66.5c 137.1b 149.1a 77.9c 15.7c 37.3a 17.0b
d
Means within a row for each constituent with the same letter are not significantly different (p > 0.05) as determined using Duncan’s multiple range test.
846 N. Wang et al. / LWT - Food Science and Technology 42 (2009) 842–848
Table 4
Effect of cooking and dehulling on minerals of lentils.
Variety Ca (mg/100 g dry matter) Cu (mg/100 g dry matter) Fe (mg/100 g dry matter)
Laird 943.0b 567.0c 1008.5a 109.0a 103.5b 83.5c 1.32b 1.42a 1.04c
Sovereign 851.0b 538.0c 943.5a 105.0a 100.1b 90.5c 1.36b 1.47a 1.19c
Richlea 885.0b 512.0c 951.0a 108.5a 94.5b 89.7c 1.35b 1.41a 1.25c
Vantage 861.5b 531.5c 914.0a 108.5a 96.3b 96.3b 1.50b 1.57a 1.31c
Eston 848.5b 542.0c 894.0a 102.8a 90.7b 76.2c 1.05b 1.11a 0.95c
Milestone 843.5b 456.5c 885.0a 99.3a 79.5b 71.7c 1.00b 1.06a 0.94c
Robin 880.5b 603.5c 946.5a 109.0a 99.8b 84.5c 1.41b 1.48a 1.29c
Blaze 896.5b 643.0c 939.5a 105.0a 95.2b 81.2c 1.28b 1.35a 1.10c
content in dehulled seeds were significantly (p < 0.05) higher than since the levels of some essential amino acids are markedly
those in the raw seeds. decreased (Youssef, Hamza, Abdel-Aal, Shekib, & El-Banna, 1986).
Dehulling resulted in a significant (p < 0.05) reduction in TIA for the
3.3. Effect of cooking and dehulling on anti-nutritional constituents eight lentil varieties. Data from this study indicated that cooking
reduced TIA more effectively than dehulling, confirming the finding
Trypsin inhibitor activity (TIA) in the raw lentil samples ranged of Wang et al. (2008).
from 1.91 to 2.77 mg/g sample (Table 5). The mean TIA value for the Phytic acid content varied from 6.2 to 8.8 g/kg dry matter in
eight varieties was 2.29 mg/g sample, which was in the range lentil varieties with a mean of 7.7 g/kg dry matter (Table 5), which
(1.94–3.07 mg/g) reported by Wang and Daun (2006). Analysis of was in agreement to our previous reported data (Wang & Daun,
variance showed that both variety and treatment (cooking and 2006). Analysis of variance showed that both variety and treatment
dehulling) had a significant effect on TIA (Table 1). The interactive (cooking and dehulling) had a significant effect on phytic content
effect of variety-by-cooking on TIA was also significant (p < 0.001) (Table 1). The interactive effect of variety-by-cooking on phytic acid
(Table 1). Variety-by-dehulling showed a significant (p < 0.001) content was significant (p < 0.01), but no interactive effect of
effect on TIA. A significant (p < 0.05) reduction in TIA was found for variety-by-dehulling on phytic acid was observed (Table 1). Cook-
all samples after cooking (Table 5). Cooking has been reported to be ing caused a significant (p < 0.05) reduction in phytic acid content
effective in inactivating protease inhibitors in pulses (Gatta, Pier- in the lentil varieties (Table 5). Similar results had been reported on
giovanni, Ng, Carnovale, & Perrino, 1989; Wang et al., 1997; Vidal- other pulses (Ologhobo & Fetuga, 1984; Vidal-Valverde et al., 1994;
Valverde et al., 1994). Sufficient cooking of lentils is necessary to not Wang et al., 2008). Dehulling significantly (p < 0.05) increased
only soften the lentils but also to inactivate or reduce TIA. However, phytic acid content in dehulled seeds (Table 5), indicating that
it is reported that over cooking reduces the nutritive value of pulses phytate is mainly concentrated in the cotyledons of lentils. This
Table 5
Effect of cooking and dehulling on trypsin inhibitor activity (TIA), phytic acid, tannin and sucrose content of lentils.
Variety TIA (mg/g dry matter) Phytic acid (g/kg dry matter) Tannins (g/kg dry matter) Sucrose (g/kg dry matter)
Raw Cooked Dehulled Raw Cooked Dehulled Raw Cooked Dehulled Raw Cooked Dehulled
Laird 2.24ad 0.39c 1.77b 8.8b 7.4c 9.4a 4.0a 2.3b 0.1c 22.7a 8.0c 20.9b
Sovereign 1.91a 0.43c 1.80b 8.0b 7.4c 8.6a 4.1ab 2.4b 0.1c 31.9a 15.7c 30.9b
Richlea 1.99a 0.42c 1.92b 7.0b 6.5c 7.4a 5.1a 2.7b 0.2c 20.8a 7.4c 18.6b
Vantage 2.24a 0.45c 1.83b 6.4b 6.0c 7.2a 5.1a 2.3b 0.1c 24.7a 10.1c 23.4b
Eston 2.39a 0.78c 2.01b 8.2b 7.7c 8.7a 6.1a 3.0b 0.1c 25.0a 11.6c 23.3b
Milestone 2.08a 0.56c 1.61b 6.2b 5.2c 6.6a 3.4a 3.1b 0.1c 25.6a 8.1c 24.6b
Robin 2.77a 0.59c 2.30b 8.7b 8.2c 9.3a 5.9a 3.0b 0.1c 21.5a 11.2c 20.7b
Blaze 2.72a 0.72c 2.17b 8.1b 7.5c 9.7a 4.0a 2.3b 0.2c 27.3a 14.9c 26.0b
d
Means within a row for each constituent with the same letter are not significantly different (p > 0.05) as determined using Duncan’s multiple range test.
N. Wang et al. / LWT - Food Science and Technology 42 (2009) 842–848 847
confirmed the observations of Beal and Mehta (1985) who reported anti-nutritional factors in lentils. Significant differences in proxi-
that the hull or seed coat fraction of pea contained little or no mate composition, dietary fiber, minerals, oligosaccharides, trypsin
phytate. The high level of phytic acid is of nutritional significance as inhibitor activity (TIA), phytic acid and tannin content were found
not only is the phytate phosphorus unavailable to humans, but it among lentil varieties. Cooking significantly increased crude
also lowers the availability of many other essential minerals such as protein, starch, insoluble dietary fiber (IDF), total dietary fiber
K and Zn (Reddy, Pierson, Sathe, & Salunkhe, 1989). (TDF), resistant starch (RS), Ca, Cu and Mn in lentils while reduced
The mean tannin content in raw seeds was 4.7 g/kg dry matter ash content, soluble dietary fiber (SDF), Fe, K, Mg, P, Zn, TIA, phytic
(Table 5), which was in the range reported by Vailancourt, Slinkard, acid, tannins, sucrose and oligosaccharides were observed. Dehul-
and Reichert (1986). Analysis of variance showed that both variety led seeds had significant higher crude protein, starch, RS, K, P,
and treatment (cooking and dehulling) had a significant effect on phytic acid, stachyose and verbascose content, but had lower TIA,
tannin content (Table 1). The interactive effect of variety-by-cook- tannin, SDF, IDF, TDF and Fe content than the raw lentils.
ing and variety-by-dehulling on tannin content was significant
(p < 0.001) (Table 1). Cooking significantly (p < 0.05) reduced
tannin content in the lentil varieties (Table 5). The results are in Acknowledgement
agreement with that reported on other pulses (Rao & Deosthale,
1982). Dehulling significantly (p < 0.05) reduced tannin content to We gratefully acknowledge the technical assistance of M.
a mean value of 0.13 g/kg dry matter in dehulled seeds (Table 5). Richardson and K. Jarrin of Canadian Grain Commission, Grain
This indicates that tannins are present mainly in seed coats, which Research Laboratory. We are also grateful to Pulse Canada for
was in agreement with the results reported (Singh, 1988; Wang, partially funding this project.
2008).
Sucrose and oligosaccharides (raffinose, stachyose and verbas-
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