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Plant Foods for Human Nutrition 60: 69–75 2005. C 2005 Springer Science+Business Media Inc. DOI: 10.1007/s11130-005-5102-y 69 Nutritional and Antinutritional Characteristics of Seven South Indian Wild Legumes V. V ADIVEL 1∗ K.JANARDHANAN 2 1 Indian Cardamom Research Institute Spices Board Myladumpara Kailasanadu 685 553 Kerala India 2 Seed Technology Laboratory Department of Botany Bharathiar University Coimbatore 641 046 Tamil Nadu India ∗ author for correspondence e-mail: Abstract. Seeds from seven species of wild legumes of the South India were analyzed for proximate composition mineral profiles amino acid profiles of total seed protein in vitro protein digestibility and certain antin- utritional factors to assess their potential as alternative sources of protein crops. The major findings of the study were as follows: crude protein ranged from 20.3 to 35.0 crude lipid 3.1–9.6 crude fiber 5.9–12.1 ash 2.7– 5.1 and carbohydrates 49.2–61.8. Minerals viz. sodium potassium calcium magnesium phosphorus iron copper zinc and manganese oc- curred in the range 42.9–135.9 556.2–1639.5 304.5–572.2 174.9–686.7 98.4–947.8 3.6–16.4 0.2–1.2 2.0–30.0 and 1.0–3.9 mg/100 g seed flour respectively. Profiles of amino acids of total seed proteins detected in the present study revealed that levels of valine phenylalanine tyrosine isoleucine and histidine of all the seven wild legume seed samples threo- nine of Canavalia ensiformis and C. gladiata leucine of Mucuna pruriens var. pruriens and lysine of Cassia floribunda and C. obtusifolia were found to be higher than FAO/WHO 1991 requirement pattern. The in vitro protein digestibility of the legumes under study ranged from 63.39 to 83.32. Antinutritional factors such as total free phenolics ranged from 0.41 to 5.96 tannins from 0.04 to 0.60 L-DOPA from 1.34 to 8.37 and trypsin inhibitor activity from 13.48 to 65.43 TIU/mg protein. The detected antinutritional factors probably have little nutritional significance if the seeds are properly processed. Keywords: Amino acid profiles Antinutritional factors Chemical com- position In vitro protein digestibility Wild legumes Introduction Nutrition is a most important basic need being a major determinant of health labour productivity and mental de- velopment. But in most developing countries of the world hunger and malnutrition are increasing due to population explosion shortage of fertile land and high food prices 1. Protein deficiency is widespread and has been cited as the most common form of malnutrition in developing countries 2. With a high protein content along with en- ergy values and the important vitamin and mineral content legumes have been recognised for their nutritional impor- tance. Legumes occupy the second place after cereals as sources of calories and protein in human diet. Nonetheless pulse production in India could not keep pace with popula- tion growth and consequently its per capita availability has declined from 70 g in 1956 to 34 g in 1996 3. Intensive efforts to find alternative sources of proteins from plants adapted to adverse conditions have been conducted around the world 4–8 and have met with some success. Most of the indigenous legumes in South India and in the tropics in general are relatively underutilized compared to chickpeas black gram cowpeas green gram horse gram pea pigeon pea kidney bean moth bean soybean and peanut. As part of our efforts to obtain more information about these legumes toward their wider utilization stud- ies were carried out to investigate the nutritional qual- ity of mature seeds of seven indigenous legumes namely: Canavalia ensiformis DC C. gladiata Jacq. DC Cassia floribunda Cav. C. obtusifolia L. Mucuna monosperma DC ex Wight M. pruriens var. pruriens L. DC and M. pruriens var. utilis Wall.ex Wight. Compositional knowl- edge of these materials could help in developing techno- logical processes to destroy eliminate or inactivate toxic antinutritional factors or make the plant material edible and more digestible. Materials and Method Materials All the investigated seven wild legume species were col- lected from different agroclimatic regions of South India Table 1. All seed samples were collected from tropical forests of Western Ghats. The wild legumes were identified by the methods of Sudhir et al. 9 Wilmot-Dear 10 and Singh and Premanath 11. The collected seeds were dried in open sunlight for 2–3 days. After thorough clearing and removal of broken seeds and foreign materials seeds were stored in airtight plastic containers at room temperature 25 ◦ C ± 2. The air-dried seeds approximately 50 g were powdered in a Wiley mill Scientific Equipment Delhi India to 60-mesh size and stored in screw-capped bottles at room temperature for further analysis. Proximate Composition Analysis The moisture content of the seed was estimated by tak- ing 50 transversely cut seeds at a time and the weight was taken before and after incubation in a hot-air-oven Toshniwal Brothers SR Private LTD Chennai India at 80 ◦ C for 24 h followed by cooling in a desiccator. Nitro- gen content in the powdered seed samples was estimated by the micro-kjeldahl method 12 and crude protein was calculated N × 6.25. The recommended methods of As- sociation of Official Analytical Chemists 13 were used for

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70 Table 1. Wild legumes from where collected in South India Name of the wild legume Locality District State Canavalia ensiformis Valacode Kollam Kerala C. gladiata Courtallum Tirunelveli Tamil Nadu Cassia floribunda Kargal Uttarkannada Karnataka C. obtusifolia Top Slip Coimbatore Tamil Nadu Mucuna monosperma Bogalthode Mysore Karnataka M. pruriens var. pruriens Arunooli Thrissur Kerala M. pruriens var. utilis Valanad Trivendrum Kerala the determination of ash crude lipid and crude fiber. Ash content was determined by incineration of 2 g of sample in a muffle furnace Tempo Instrument Equipment I Private LTD Bombay India kept at 600 ◦ C for 6 h. Crude lipid was determined by exhaustively extracting 2 g of sample with petroleum ether using a Soxhlet apparatus. Crude fiber was determined by acid and alkaline digestion methods. The car- bohydrate content was calculated by subtracting the total of the percentages of crude protein crude lipid crude fiber and ash on moisture-free basis from 100 14. The energy value of the seed was estimated in kJ by multiplying the percentages of crude protein crude lipid and carbohydrate by the factors 16.7 37.7 and 16.7 respectively 6. Three samples for each species were analyzed and the results expressed on dry weight basis. Mineral Analysis Five hundred mg of the ground legume seed was digested with a mixture of 10 ml concentrated nitric acid 4 ml of 60 perchloric acid and 1 ml of concentrated sulphuric acid. After cooling the digest was diluted with 50 ml deion- ized distilled water filtered with Whatman No. 42 filter paper and filtrates made up to 100 ml in a glass volumetric flask with deionized distilled water. All the minerals except phosphorus were analyzed from triple acid digested sample by an atomic absorption spectrophotometer Model 5000 Perkin-Elmer Boston MA USA equipped with different lamps 15. Phosphorus content in the triple acid digested extract was colorimetrically analyzed 16 at 650 nm using a spectrophotometer Model Spectronic 20D Milton Roy Ivyland PA USA. Amino Acid Analysis Total protein was extracted by a modified method of Basha et al. 17. The ethanol treatment was omitted to save the prolamin fraction. The extracted proteins were puri- fied by precipitation with cold 20 w/v trichloroacetic acid TCA and estimated by the method of Lowry et al. 18. A 30 mg protein sample was hydrolyzed by 6 N HCl 5 ml in an evacuated sealed glass tube which was kept in air oven Toshniwal Brothers SR Private LTD Chennai India maintained at 110 ◦ C for 24 h. The sealed tube was broken and the acid removed completely by repeated flash evaporation after the addition of deionized water. Dilution waseffected by means of citrate buffer pH 2.2 to such an ex- tent that the solution contained 0.5 mg protein/ml. The solu- tion was filtered through a Millipore filter 0.45 µm Waters Millipore Mississauga ON Canada prior to injection. Us- ing precolumn derivation with O-phthaldialdehyde OPA amino acid analysis was performed. The unit was a reversed- phase HPLC Model 23250 ISCO Lincoln NE USA fit- ted with spherisorp C 18 column 4.6mm × 250 mm and ISCO-dual pump. The flow rate was 1.5 ml/min and a flu- orescence detector excitation 305–395 nm emission 430– 470 nm was used. The contents of the different amino acids recovered are expressed as g/100 g protein. The essential amino acid score was calculated as follows: Essential amino acid score grams essential amino in 100 g of test protein grams essential amino in 100 g ofFAO/WHO 19reference pattern × 100 Determination of In Vitro Protein Digestibility IVPD Protein digestibility was assayed by the in vitro method de- scribed by Hsu et al. 20. The enzymes used for IVPD were purchased from Sigma Chemical Co. St. Louis MO USA. Calculated control casein and samples were weighed out hydrated in 10 ml of distilled water and refrigerated at 5 ◦ C for 1 h. The protein-containing samples and enzymes were all adjusted to pH 8.0 at 37 ◦ C. The IVPD was determined by the sequential digestion of the protein-containing sample with a multienzyme mixture trypsin porcine pancreatic trypsin—Type IX with 14190 BAEE unites/mg protein α-chymotrypsin bovine pancreatic chymotrypsin—Type II 60 unites/mg powder and peptidase porcine intestinal peptidase—Grade III 40 units/g powder at 37 ◦ C followed by protease Type IV from Streptomyces griseusat55 ◦ C. The pH drop of the samples from pH 8.0 was recorded after

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71 20 min of incubation. The IVPD was calculated according to the regression equation Y 234.84–22.56 X where Y digestibility X pH drop described by Hsu et al. 20. Analysis of Antinutritional Compounds The antinutritional compounds total free phenolics 21 tannins 22 and the nonprotein amino acid L-DOPA 34- dihydroxyphenylalanine 23 were quantified. Trypsin inhibitor activity was determined by the enzyme assay of Kakade et al. 24 by using benzoyl-DL-arginine-p- nitroanilide BAPNA as a substrate. One trypsin inhibitor unit TIU has been expressed as an increase of 0.01 ab- sorbance units per 10 ml of reaction mixture at 410 nm. Trypsin inhibitor activity has been defined in terms of trypsin units inhibited per mg protein. Results and Discussion The results of the proximate composition of the seven wild legumes are presented in Table 2. The crude protein val- ues ranged between 20.3 in Cassia obtusifolia and 35 in Canavalia ensiformis. The protein values are compa- rable to those reported for some cultivated legume seeds 25 like Bengal gram 17.1 cowpea 24.1 filed bean 24.9 green gram 24.0 lentil 25.1 moth beans 23.6 and peas 19.7 but are lower than the soybean 43.2. These legumes are used extensively in typical In- dian diets and are expected to play a significant role in improving protein nutrition in India. The remarkably high level of protein in the wild legumes under study underscores their importance as a potential source of this vital nutrient. The crude fat contents of the seeds varied between 3.1 in Cassia floribunda and 9.6 in Mucuna monosperma. The crude fat content does not qualify the presently investigated wild legumes as oil rich legumes especially when compared with groundnut and soybeans which have fat contents of about 25.3 and 19.5 respectively 25. Relative to the levels reported for Indian edible legumes such as Bengal Table 2. Proximate composition of the wild legumes collected from South India g/100 g seed flour a Energy Sample Moisture Crude protein Crude lipid Crude fiber Ash CHO b kJ/100 gDM Canavalia ensiformis 8.5 ± 0.3 35.0 ± 0.8 4.3 ± 0.5 7.7 ± 0.3 3.9 ± 0.1 49.2 1568 C. gladiata 8.5 ± 0.5 25.5 ± 0.6 3.3 ± 0.4 5.9 ± 0.2 3.5 ± 0.1 61.8 1582 Cassia floribunda 5.7 ± 0.1 21.7 ± 0.6 3.1 ± 0.3 10.8 ± 0.9 3.4 ± 0.1 61.0 1497 C. obtusifolia 6.8 ± 0.2 20.3 ± 0.3 7.4 ± 0.2 6.8 ± 0.1 5.1 ± 0.1 60.3 1626 Mucuna monosperma 6.5 ± 0.1 21.2 ± 0.6 9.6 ± 0.1 12.1 ± 0.1 2.7 ± 0.1 54.4 1625 M. pruriens var. pruriens 8.4 ± 0.2 32.4 ± 0.6 5.7 ± 0.4 7.8 ± 0.3 3.6 ± 0.2 50.6 1601 M. pruriens var. utilis 8.5 ± 0.2 29.3 ± 1.2 6.4 ± 0.5 8.4 ± 0.3 4.2 ± 0.1 51.7 1593 a All values are mean of triplicate determination expressed on dry weight basis ± standard error. b CHO—Carbohydrates calculated by difference. gram cowpeas green gram horse gram moth beans and peas 25 a high range 5.9–12.1 of fiber is recorded in the presently investigated wild leguminous seeds. The World Health Organization WHO has recommended an intake of 22–23 g of fiber for every 1000 kcal of diet 26. Though it does not contribute to the nutritive value of foods the presence of fiber i.e. roughage in the diet is neces- sary for digestion and for efficient elimination of wastes. The contraction of muscular walls of the digestive tract is stimulated by fiber thus counteracting constipation 25. Therefore the high range of fiber recorded in the presently investigated wild legumes is a desirable characteristic since fiber plays an important role in the diet of humans and an- imals. The ash content is within 2.7–5.1. This range is similar to that found in the literature for legumes that serve as food sources of minerals. It appears that the presently investigated wild legumes have a high range of carbohydrate 49.2–61.8 because of their low fat content. Groundnut and soybeans have lower carbohydrate values of 26.1 and 20.9 respectively 25. All seven wild legumes exhibit higher energy values 1497–1626 kJ/100 g DM than con- ventional pulse crops like cowpea green gram horse gram moth bean and peas 25 which are in the range of 1318– 1394 kJ/100 g DM. The data on mineral profiles are given in Table 3. Potas- sium as in most legumes is the predominant mineral ex- cept in Cassia floribunda where phosphorus is predominant. The seeds of Cassia obtusifolia exhibit the highest level of calcium 572.2 mg/100 g magnesium 686.7 mg/100 g phosphorus 947.8 mg/100 g and zinc 30 mg/100 g while the seeds of Mucuna pruriens var. pruriens are rich in iron 16.4 mg/100 g. The seeds of M. pruriens var. utilis contain the highest levels of copper and manganese. In general the contents of sodium calcium magnesium iron copper and manganese are found to be high compared to reported levels for Pisum sativum 27. Table 4 shows the amino acid composition of total seed proteins and the essential amino acid scores of wild legumes along with FAO/WHO 19 reference pattern. The data on the amino acid pattern of total seed proteins of

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72 Table 3. Mineral constituents in wild legumes collected from South India mg/100 g seed flour a Sample Na K Ca Mg P Fe Cu Zn Mn Canavalia ensiformis 56.8 ± 3.8 1017.5 ± 2.8 497.9 ± 9.1 192.1 ± 7.5 240.2 ± 5.2 5.2 ± 3.2 0.4 ± 0.9 4.3 ± 1.7 1.0 ± 0.2 C. gladiata 109.3 ± 2.7 1639.5 ± 3.4 510.1 ± 6.2 480.9 ± 4.6 601.2 ± 3.5 10.9 ± 0.9 0.8 ± 0.2 6.6 ± 0.2 2.2 ± 0.1 Cassia floribunda 135.9 ± 6.4 573.2 ± 8.4 537.8 ± 3.5 288.7 ± 5.4 603.1 ± 6.7 3.6 ± 1.0 0.3 ± 1.0 2.0 ± 0.5 1.0 ± 0.5 C. obtusifolia 42.9 ± 7.3 1555.8 ± 3.8 572.2 ± 4.4 686.7 ± 5.8 947.8 ± 6.4 10.7 ± 0.7 1.0 ± 0.8 30.0 ± 0.6 2.1 ± 0.5 Mucuna monosperma 98.4 ± 4.4 556.2 ± 5.6 313.8 ± 3.7 273.8 ± 8.5 206.8 ± 7.3 4.8 ± 0.9 0.2 ± 0.4 7.7 ± 0.5 1.1 ± 0.2 M. pruriens var. pruriens 47.8 ± 6.8 835.1 ± 5.7 304.5 ± 7.4 208.8 ± 4.4 119.6 ± 3.8 16.4 ± 0.4 1.0 ± 0.8 2.3 ± 0.5 1.8 ± 0.4 M. pruriens var. utilis 52.5 ± 1.2 778.1 ± 1.4 393.4 ± 7.5 174.9 ± 4.4 98.4 ± 3.5 13.4 ± 1.2 1.2 ± 0.6 6.7 ± 0.4 3.9 ± 0.2 a All values are mean of triplicate determination expressed on dry weight basis ± standard error. the investigated wild legumes reveal that threonine con- tent of Cassia floribunda C. obtusifolia Mucuna pruriens var. pruriens and M. pruriens var. utilis leucine content of Canavalia ensiformis C. gladiata Cassia floribunda Mucuna monosperma M. pruriens var. utilis and lysine content of Canavalia gladiata Mucuna monosperma M. pruriens var. pruriens and M. pruriens var. utilis seem to be deficient whereas the contents of valine phenylalanine tyrosine isoleucine and histidine of all the seven wild legume seed samples threonine content of Canavalia ensi- formis and C. gladiata leucine content of Mucuna pruriens var. pruriens and lysine content of Cassia floribunda and C. obtusifolia are found to be higher than FAO/WHO 19 requirement pattern. In general the amino acid profiles are an underestimate of total amino acid levels because some amino acids are destroyed during the preparation of the sample by acid digestion. The in vitro protein digestibility IVPD of all seven wild legumes studied ranged from 63.39 to 83.32 Table 5. This range seems to be higher compared to pigeon pea 28 29 and green gram 30 and is comparable with that of cow- pea 7 31 32 chickpea 31 33 34 horse gram 31 35 kidney bean 36 moth bean 35 and soybean 33. The highest IVPD is recorded for Cassia floribunda followed by Mucuna monosperma.However these values are much lower than the corresponding value for casein 97.7 37. The presence of antinutritional factors is one of the major drawbacks limiting the nutritional and food qualities of the legumes 38. A preliminary evaluation of some of these fac- tors in raw wild legumes is made for this reason Table 5. In the studied seeds total free phenolics occurred within the range of 0.41–5.96 and tannins ranged from 0.04 to 0.60. These ranges seem to be lower compared to black gram 39. The negative nutritional effects of tannins are diverse and incompletely understood but the major effect is to cause growth depression by decreasing the digestibility of protein and carbohydrate. This is most likely the conse- quence of the interaction of tannins with either protein or starch to form enzyme-resistant substances 38. Tannins are concentrated mainly in the seed coat 5 8 preliminary dehulling constitutes the simplest method for their removal. Available information also indicates that the levels of tan- nin in legumes can be reduced by simple processing meth- ods such as soaking roasting and autoclaving 6 39–41. The L-DOPA content in both species of Canavalia ranged from 2.64 to 2.83 of dry weight. In Cassia the L-DOPA content ranged from 1.34 to 1.57 and in Mucuna from 4.52 to 8.37. In M. pruriens var. pruriens it accounts for 8.37. In earlier studies it has been demonstrated that the level of L-DOPA is significantly reduced by repeated soak- ing and boiling of seeds 42 dry heat 6 and autoclaving 41. Trypsin inhibitor activity ranged from 13.48 TIU/mg proteins in Cassia obtusifolia to 65.43 TIU/mg proteins in Mucuna monosperma. All legumes studied to date have been found to contain trypsin inhibitors in varying amounts. Though the trypsin inhibitor activity has been studied in a number of pulses the results obtained in the present investigation cannot be compared because the expression of trypsin inhibitor activity nature and concentration of the substrate etc. are different. However based on the in- vestigations carried out under similar experimental condi- tions for trypsin inhibitor activity in cultivated legumes like pigeon pea 67.1–71.3 TIU/mg proteins 28 and kidney bean 79.7–87.6 TIU/mg proteins 43 it may be inferred that the trypsin inhibitor activity obtained in the present in- vestigation appears to be low. Trypsin inhibitors ingested in significant amounts disrupt the digestive process and may lead to undesirable physiological reactions. Trypsin inhibitor is thermolabile and its inhibitory activity can be reduced considerably by thermal treatment 38. In conclusion this study reveals that the chemical com- position of all seven wild legumes seem to be compara- ble to that of cultivated legumes reported earlier. In ad- dition they contain higher levels of certain amino acids compared to recommended levels. In vitro protein di- gestibility of these legumes is also found to be higher than that of certain common legumes. The presence of antinu- tritional factors identified in this current report should not pose a problem to human health if the seeds are properly processed. In view of the overall nutrient and chemical composition seeds of these wild legumes can be explored as an alternative protein source to alleviate the protein- energy-malnutrition that can occur among economically weaker segments of the population in developing countries.

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73 Table 4. Amino acid profiles of acid hydrolyzed purified total seed proteins g/100 g proteins Amino Canavalia Cassia Mucuna M. pruriens M. pruriens FAO/WHO 19 acid ensiformis EAAS C. gladiata EAAS floribunda EAAS C. obtusifolia EAAS monosperma var. pruriens EAAS var. utilis EAAS requirement pattern Asp 12.53 14.02 10.83 13.52 11.23 10.53 17.14 Glu 16.38 19.37 18.82 20.71 16.85 9.84 9.56 Ala 3.12 5.12 3.27 3.84 6.93 2.55 3.87 Val 4.83 138 4.03 115 4.38 125 4.18 119 4.93 141 8.57 245 7.35 210 3.5 Gly 5.24 5.25 5.74 5.46 4.43 10.73 5.12 Arg 7.83 4.84 6.85 6.47 2.38 7.14 5.14 Ser 6.54 3.73 4.04 4.82 3.84 5.75 4.93 Cys N.D N.D N.D N.D N.D N.D N.D 2.5 Met 1.78 1.51 0.83 0.99 0.34 1.27 1.24 Thr 4.03 118 4.02 118 2.87 84 3.21 94 3.57 105 2.58 76 3.27 96 3.4 Phe 4.83 5.23 125 5.82 122 4.40 108 5.87 172 3.38 137 3.78 137 6.3 Tyr 3.45 131 2.62 1.87 2.43 4.95 5.27 4.88 Ile 2.87 102 4.44 158 3.83 137 4.19 149 5.87 210 3.96 141 2.94 105 2.8 Leu 3.95 60 4.03 61 6.23 94 6.61 100 4.97 73 7.28 110 6.34 96 6.6 His 3.73 196 3.12 164 5.27 277 2.08 109 2.38 125 4.27 225 4.17 219 1.9 Lys 5.78 100 4.85 83 6.47 115 7.16 123 5.48 94 5.16 89 4.95 85 5.8 Try N.D. N.D. N.D. N.D. N.D. N.D. N.D 1.1 Pro N.D. N.D. N.D. N.D. N.D. N.D. N.D Note. The amino acid profiles are an underestimate of total amino acid levels because the amino acids cystine and tryptophan are destroyed during the preparation of the sample by acid digestion. N.D.—Not Detected EAAS—Essential amino acid score.

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74 Table 5. In vitro protein digestibility IVPD and antinutritional factors in wild legumes collected from South India Trypsin inhibitor Activity a Sample IVPD a Total free phenolics b Tannins b L-DOPA b TIU/mg protein Canavalia ensiformis 74.66 1.23 ± 0.05 0.16 ± 0.01 2.64 ± 1.31 34.34 C. gladiata 63.39 1.94 ± 0.13 0.20 ± 0.02 2.83 ± 0.05 26.83 Cassia floribunda 83.32 0.41 ± 0.52 0.42 ± 0.12 1.57 ± 0.23 16.84 C. obtusifolia 74.66 0.66 ± 0.05 0.60 ± 0.05 1.34 ± 0.45 13.48 Mucuna monosperma 78.32 2.29 ± 0.68 0.55 ± 0.21 4.52 ± 0.92 65.43 M. pruriens var. pruriens 72.41 5.96 ± 0.11 0.05 ± 0.01 8.37 ± 0.39 62.34 M. pruriens var. utilis 72.41 4.48 ± 0.13 0.04 ± 0.43 6.68 ± 0.07 48.23 a Values of two independent experiments. b Values are mean of triplicate determination expressed on dry weight basis ± standard error. Acknowledgment We thank the Government of India Ministry of Environ- ment Forests New Delhi India for financial assistance. References 1. 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