Food Bioscience 34 (2020) 100527

Contents lists available at ScienceDirect

Food Bioscience
journal homepage: www.elsevier.com/locate/fbio

Nutritional, anti-nutritional and physico-functional properties of wild edible T

yam (Dioscorea spp.) tubers from Koraput, India
Bandana Padhan, Meghali Biswas, Debabrata Panda∗
Department of Biodiversity and Conservation of Natural Resources, Central University of Orissa, Koraput, 764021, Odisha, India

A R T I C LE I N FO A B S T R A C T

Keywords: Wild yams (Dioscorea spp.) make a significant contribution to the diets and economic welfare of tribal people of
anti-nutritional factors Koraput, India. However, there is a lack of information on the food quality of the tubers. The present study was
Dioscorea carried out to evaluate the proximate, nutritional and anti-nutritional compositions as well as the physico-
Wild yam functional properties in 8 wild and one cultivated Dioscorea species. The proximate compositions, i.e., moisture,
ash, fat, fiber, and crude protein content of yam tubers ranged from 54 to 83, 2.5 to 5.4, 0.56 to 2.0, 1 to 2.02
and 5.4–10.3% respectively. The mineral composition ranged from 31.4 to 89.4 mg 100 g−1 of sodium,
0.83–1.36 g 100 g−1 of potassium, with significant amounts of vitamin C and E. In addition, some wild species
showed better nutritional and physico-functional parameters in comparison to the cultivated ones. Furthermore,
the anti-nutritional compounds such as diosgenin, amylase- and trypsin-inhibitors were also significantly higher
in some wild Dioscorea species. The biochemical characterization with multivariate data analysis led to a feasible
classification of the usage of wild yam species. Taken together, the wild species, such as D. oppositifolia, D.
hamiltonii and D. pubera showed better nutritional composition than the other yam species. The study also
suggested that these wild yam species are a safe food source for local consumption and domestication, leading to
potential improvement of food security.

1. Introduction provide dietary energy from carbohydrates in developing countries

(Ofosu, 2012, pp. 1–4; Otegbayo et al., 2018). There is an enormous
Food and nutritional security are major concerns in many countries. diversity in the wild and domesticated species that are being used by
To diversify present-day agriculture as well as to search for alternative tribal communities as traditional food (Adepoju, Boyejo, & Adeniji,
food and feed ingredients has the potential to address food insecurity 2018; Behera et al., 2009). However, systematic characterization of
(Adebooye & Phillips, 2006; Otegbayo, Oguniyan, Olunlade, Oroniran, food quality traits in wild species is a major prerequisite for greater
& Atobatele, 2018). Some wild root and tuber crops are important for consumption and possible cultivation. Food quality in yam tubers in-
food security of the developing world due to their high caloric value cludes proximate, nutritional and anti-nutritional analysis (Otegbayo
and carbohydrate content (Bhandari, Kasai, & Kawabata, 2003; Price, et al., 2018).
Bhattacharjee, Montes, & Fraser, 2017). Yams are the staple food of Koraput is one of the tribal dominated districts of Odisha in India
millions of people especially in the tropics and sub-tropics and have a (18° 14ʹ to 19° 14ʹ N latitude and 82° 05ʹ to 83° 25ʹ E longitude) and was
significant place in the dietary habits of many rural and tribal com- recently declared as one of the agro-biodiversity hot spots in India
munities (Behera, Maharana, Sahoo, & Prusti, 2009; Bhandari et al., (Mishra & Chaudhury, 2012). There are 8 wild Dioscorea species, i.e.,
2003). Dioscorea species are monocotyledonous tuber crops of the fa- Dioscorea oppositifolia L., D. hamiltonii Hook.f., D. bulbifera L., D. pubera
mily Dioscoreacea and the genus includes more than 600 different Blume., D. pentaphylla L., D. wallichii Hook.f., D. glabra Roxb. and D.
species (Amanze, Agbo, Eke-Okoro, & Njoku, 2011; Kouakou, Dabonne, hispida Dennst. along with one cultivated species D. alata L., which have
Guehi, & Kuoame, 2010). A few species are commercially cultivated been used as food by the tribal people (Mishra, Swain, Chaudhary, &
and some wild edible species have not yet been domesticated due to Ray, 2011; Padhan & Panda, 2016). Yam tubers are diverse and they are
inferior tuber quality, low yield and the presence of anti-nutritional consumed in different forms, mostly boiled, fried, or baked. The che-
compounds (Behera et al., 2009). Dioscorea species have the potential to mical and nutritional composition of the Dioscorea species of Nepal

∗
Corresponding author. Department of Biodiversity and Conservation of Natural Resources, Central University of Orissa, Landiguda, Koraput, Odisha, 764021,
India.
E-mail address: [email protected] (D. Panda).

https://doi.org/10.1016/j.fbio.2020.100527
Received 17 March 2018; Received in revised form 5 January 2020; Accepted 5 January 2020
Available online 07 January 2020
2212-4292/ © 2020 Elsevier Ltd. All rights reserved.
B. Padhan, et al. Food Bioscience 34 (2020) 100527

Fig. 1. Tubers of different wild and cultivated Dioscorea species used by the tribal people of Koraput, India.

(Bhandari et al., 2003; Bhandari & Kawabata, 2004), India (Mohan & Chennai, India) for 3 h using petroleum ether (100 ml) as solvent and
Kalidass, 2010; Shajeela, Mohan, Jesudas, & Soris, 2011; the difference in the weight of the solvent beaker was considered as the
Shanthakumari, Mohan, & Britto, 2008), Ghana (Polycarp, Afoakwa, fat content of the yam (AOAC, 2012-920.39).
Budu, & Otoo, 2012), Nigeria (Otegbayo et al., 2018) and China (Wu The ash content was determined by taking dry sample (2 g) and
et al., 2016) have been studied. No data on the chemical compositions burning at 600 °C for 3 h in a muffle furnace using the AOAC,
of wild edible Dioscorea species was found in the literature. The present 2012–923.03 method. The weight of the residue was measured as ash
study was undertaken to determine the nutritional compositions, anti- content and expressed as a percentage.
nutritional factors and physico-functional properties of different wild The moisture and fat free samples (2 g) were used for crude fiber
Dioscorea tubers of Koraput. determination using a H2SO4 and NaOH (1.25%) extraction using the
AOAC, 2012–978.10 method. After both extractions, the precipitate
2. Materials and methods was rinsed with 10% HCl and absolute ethanol (HiMedia, Mumbai,
India) and allowed to dry overnight at 105 °C. The weight of the residue
2.1. Collection and preparation of sample was taken. The sample was heated at 600 °C for 90 min in a muffle
furnace and weight of the residue was expressed as a percentage of
The study was carried out with the 8 wild edible yam tubers along crude fiber.
with the one local cultivated species (Fig. 1). The mature tubers of The crude protein was determined using Kjeldahl method (AOAC,
different species were collected from the medicinal plant garden of the 2012-981.10). The dried yam samples were digested in 15 ml of con-
Department of Biodiversity and Conservation of Natural Resources, centrated H2SO4 and 3 g of catalyst mixture of K2SO4:CuSO4 (10:1)
Central University of Orissa, Koraput, India, which were all grown with with gentle swirling. The digested sample was made up the volume to
the same climatic and agronomic condition. Briefly, uniform sized tu- 100 ml and distillation was performed in a distillation system (KEL
bers of each species were directly sown in 30 kg polythene bags filled PLUS-DISTYL EM, Chennai, India). The digested sample of 10 ml was
with a mixture of farm soil collected from the experimental garden of mixed with equal volume of 0.5 N NaOH. Distillation was continued for
Central University of Orissa and farmyard manure (cow dung) with a 10 min and the distillate was collected in a separate conical flask
3:1 ratio. The plants were regularly irrigated with tap water and sub- containing 20 ml of 4% boric acid solution with 2–3 drops of methyl red
jected to natural solar radiation. Harvesting of tubers was done after all and bromocresol green indicator. The distillate was titrated against
the vines dried which was around 240 days after planting. After col- standard 0.1 N HCl solutions until the appearance of a pink colour
lection, the tubers were washed, peeled, cut into slices of ~2.5 mm against a blank. The percentage of crude protein was calculated using
diameter using a knife and shade dried at 25 ± 2 °C for 4–6 days. The the following formula:
dried tuber samples were mechanically ground in a pulverizer (SK-03, Crude protein (%) = 6.25 × % N
Global Instrumentation, Visakhapatnam, India) into powder and sieved
(2 mm particle size) and stored in airtight containers for a maximum of
8 wk. 2.3. Nutritional composition

2.2. Proximate composition Carbohydrate (sugar and starch) content was estimated for the dry
powdered samples using the procedure of Sadasivam and Manickam
The moisture content was determined by drying the fresh tuber in (2007). Briefly, the yam sample (0.1 g) was extracted with 10 ml 80%
an oven at 100 ± 5 °C for 24–48 h to a constant weight and expressed methanol and the mixture was centrifuged at 1.118 × g 10−4
as a percentage (AOAC, 2012-934.06). (10000 rpm) (R-1500, Remi Laboratory Instruments, Mumbai, India)
The crude fat contents of the samples (1 g) were determined by for 15 min. The extraction was repeated twice and the pooled super-
continuous extraction in a lipid extractor (SOCS PLUS SCS-03E, Pelican, natant was evaporated until the methanol was removed. The extract

2
B. Padhan, et al. Food Bioscience 34 (2020) 100527

was brought to 25 ml with distilled water. The extract was used for distilled water, heated for 20 min and centrifuged at 4.1 × 10−4 x g
sugar estimation. The remaining residues were dried and re-extracted (6000 rpm) for 15 min. An aliquot of 0.5 ml extract was diluted to 10 ml
with 5 ml of 9.2 N perchloric acid followed by 5 ml of 4.6 N perchloric and mixed with 0.5 ml Folin Dennis reagent (100 g sodium tungstate,
acid for starch estimation. The sugar and starch extract (1 ml) was 2 g phosphomolybdic acid with 50 ml phosphoric acid) and followed by
added to anthrone reagent (4 ml) (0.2% in concentrated H2SO4) addition of 1 ml 10% Na2CO3 solution and incubated for 30 min. The
(Merck, Mumbai, India) and the absorbance was measured at 630 nm in intensity of colour was measured at 760 nm and the tannin content was
a spectrophotometer (Systronics-104, Systronics, Ahmedabad, India). expressed as mg g−1 dw using tannic acid (HiMedia) as a standard.
The value was expressed as mg g−1 dw using glucose and potato soluble Total alkaloid content was estimated using the gravimetric method
starch (Merck) as standards. of Harborne (1973). The yam samples (1 g) were extracted with 40 ml
The amylose content was estimated using the procedure of 10% acetic acid in methanol and incubated for 4 h. The solution was
Sadasivam and Manickam (2007), where the yam samples (100 mg) filtered and the filtrate was evaporated in a water bath at 80–90 °C. To
were extracted in a mixture of 80% ethanol and 1 N NaOH solution the extract, ammonium hydroxide solution was added dropwise
(1:5). An aliquot was prepared with 2.5 ml diluted extract, 3 drops through a dropper until the precipitation was complete. The solution
0.1% phenolphthalein (in 95% ethanol) and 0.1 N HCl was added to the was incubated for 2–3 h and the precipitate was filtered and the filtrate
mixture followed by colour development after addition of 1 ml iodine was washed with 1% ammonia solution and allowed to stand for
reagent (1 g I2 and 10 g KI in 500 ml distilled water) and the colour 30 min. The residue was oven dried and weighed to quantify the al-
change was measured at 590 nm. The amylose content from the tuber kaloid and the value was calculated as mg 100 g−1 dw.
was expressed as mg g−1 dw using amylose from potato (Sigma-Aldrich The saponin content in the yam samples was determined using the
Co., St. Louis, MO, USA) as a standard. method of Nahapetian and Bassiri (1975). The samples were extracted
The ascorbic acid (vitamin C) content was estimated using the in 20% ethanol and re-extracted with diethyl ether and n-butanol in a
method of Omaye, Turnbull, and Sauberlich (1979) with some mod- separating funnel. The mixture was washed twice with 5% aqueous
ification. The sample (1 g) was extracted with 4 ml of 10% TCA. The sodium chloride. The upper layer was collected and evaporated. After
extract of 0.5 ml was mixed with 0.1 ml DTC reagent (0.4% thiourea, evaporation the samples were dried in the oven to constant weight. The
0.05% copper sulphate and 3% 2, 4-dinitrophenylhydrazine (DNPH) in saponin content was calculated as mg g−1 dw.
9 N H2SO4) followed by incubation at 37 °C for 3 h. Chilled 65% H2SO4 The α-amylase inhibitor was evaluated using the spectro-
(0.75 ml) was added to the mixture and incubated for 30 min at room photometric method of Alonso, Orúe, and Marzo (1998). One unit of
temperature (25 ± 2 °C). The colour intensity was measured at enzyme activity was determined from soluble starch and expressed as
520 nm against a blank containing 10% TCA. Values were expressed as 1 μmol of reducing groups (calculated as maltose) min−1 at 37 °C using
mg ascorbic acid 100 g−1 dw using a known concentration of ascorbic the specified conditions. The absorbance was measured at 540 nm. One
acid (HiMedia) for the calibration. inhibitory unit of α-amylase activity was determined as one α-amylase
Vitamin E (α-tocopherol) content was estimated using the method inhibitory unit (IU) and expressed as IU g−1 dw.
of Baker, Frank, Angelis, and Feingold (1980). Powdered sample (1.0 g) Trypsin inhibitor activity (TIA) was determined using the method of
was extracted with petroleum ether (2 ml) and absolute ethanol Smith, Megen, Twaalfhoven, and Hitchcock (1980). The yam sample
(1.6 ml) and the reaction was initiated with the addition of 0.2 ml of (1 g) was extracted with 50 ml of 10 mM NaOH and was vigorously
0.2% 2,2′-dipyridyl (HiMedia) in absolute ethanol and left in the dark stirred for 1 h and centrifuged at 1.118 × 10−4 x g (10000 rpm) for
for 5 min. The colour in the aqueous layer was measured at 520 nm. 10 min. An aliquot was prepared with 1 ml sample extract, 1 ml dis-
The values are expressed as mg vitamin E 100 g−1 dw using a cali- tilled water and 2 ml standard trypsin solution. The mixture was in-
bration curve of α-tocopherol (HiMedia) as a standard. cubated at 37 °C for 10 min and 5 ml of BPNA, benzoyl-DL-arginine-p-
Minerals, such as Na and K were determined by digesting the sample nitroanilide hydrochloride (HiMedia) solution was added. The absor-
(1 g each) with a mixture of nitric and perchloric acid (9:4) at 100 °C bance of the supernatant was measured at 410 nm. TIA value was ex-
using the AOAC, 2012, p. 956.01 method. The digested sample was pressed as weight of trypsin inhibited g−1 dry matter and calculated
diluted to the suitable concentration and aspirated into a Systronics- using the formula:
105 flame photometer (Systronics, Ltd.) using analytical grade KCl
TIA= (2.632 × Dilution factor × Ai) / S mg pure trypsin inhibited g−1
(HiMedia) and NaCl (HiMedia) as standards.
sample
2.4. Anti-nutritional composition Where Ai is the change in absorbance due to trypsin inhibition ml−1
extract and S is the sample weight.
Diosgenin content was estimated using a spectrophotometric
method as described by Uematsu, Hirata, Saito, and Kudo (2000) where
the tuber sample was extracted in methanol and the extract was reacted 2.5. Physico-functional properties
with the reagent containing 0.5% p-anisaldehyde and 50% H2SO4 in
ethyl acetate. The absorbance of the coloured solution was measured at The water absorption capacity (WAC) and water solubility index
430 nm using diosgenin (HiMedia) as the standard and the value was (WSI) of Dioscorea tubers were evaluated using the method Phillips,
expressed as mg g−1 dw diosgenin equivalence. Chinnan, Branch, Miller, and Mcwatters (1988) and Anderson, Conway,
The phenol content was determined using the method of Sadasivam Pfeifer, and Griffin (1969), respectively. The tuber flours (2.5 g) were
and Manickam (2007) with some modifications. Tuber samples (0.1 g) weighed into a centrifuge tube (Mo) and 30 ml distilled water was
were refluxed in 80% methanol and evaporated and the volume of the added. The content of the centrifuge tube was shaken for 30 min. The
crude extract was made up to 10 ml. An aliquot was prepared with 1 ml mixture was kept in a water bath (37 °C) for 30 min and centrifuged at
extract, 0.5 ml Folin-Ciocalteu's reagent (Himedia) (2:1) and 2 ml 20% 5000 x g for 15 min. The resulting sediment (M2) was weighed and then
Na2CO3 and the mixture was incubated in a water bath at 90–95 °C for dried at 105 °C to constant weight (M1). The WAC was then calculated
2 min. The absorbance was measured at 650 nm and the total phenol as follows:
content was expressed as gallic acid equivalence (GAE) mg g−1 dw WAC (%) = [(M2 - M1)/M1] × 100
using GA (HiMedia) as a standard.
Total tannin content of yam flour was determined using the Folin- While the WSI was calculated using the equation:
Dennis spectrophotometric procedure described by Sadasivam and
WSI (%) = [(Mo - M1)/Mo] × 100
Manickam (2007). The tuber samples (0.1 g) were dissolved in 20 ml

3
B. Padhan, et al. Food Bioscience 34 (2020) 100527

Table 1
Variation in proximate and physico-functional parameters in different wild and cultivated species of Dioscorea. Data are the mean of three replications ± SD. Means
followed by a common letter in the same column are not significantly different at the 5% level by Fisher's least significance difference (LSD) test. WAC: water
absorption capacity; FC: foam capacity; PC: paste clarity; WSI: water solubility index; IAS: iodine affinity of starch; CV: coefficient of variance.
Species Moisture Ash content (%) Crude fat (%) Crude fibre (%) Crude protein WAC (%) FC (%) PC (%) WSI (%) IAS (ppm)
content (%) (%)

D. oppositifolia 66 ± 1d 4.4 ± 0.3b 1.62 ± 0.03b 1.52 ± 0.01b 9.5 ± 0.2b 2.2 ± 0.0b 20 ± 1b 38 ± 1a 23 ± 0.1b 4.0 ± 0.1c
D. hamiltonii 77 ± 1b 5.4 ± 0.4a 2.0 ± 0.1a 1.5 ± 0.1b 10 ± 0.1b 2.4 ± 0.0a 18 ± 1b 37 ± 0.2a 20.3 ± 0.2c 3.7 ± 0.1d
D. pubera 74 ± 1bc 4.2 ± 0.5b 1.55 ± 0.03b 1.6 ± 0.1b 10.3 ± 0.1a 2.2 ± 0.0b 22 ± 2a 38 ± 1a 25 ± 1a 5.3 ± 0.1a
D. wallichii 63 ± 0d 4 ± 0.2c 0.91 ± 0.16c 2.02 ± 0.01a 8.4 ± 0.3d 1.6 ± 0.0e 16.2 ± 0.3b 24 ± 2c 21.3 ± 0.3bc 4.2 ± 0.0b
D. hispida 79 ± 1b 2.7 ± 0.3d 0.57 ± 0.11d 1 ± 0d 5.4 ± 0.1e 1.4 ± 0.0e 13 ± 1c 17 ± 1d 13.2 ± 0.5e 2.6 ± 0.1f
D. pentaphylla 83 ± 1a 2.6 ± 0.3d 0.6 ± 0.1d 1.3 ± 0.1c 9.2 ± 0.2c 2.0 ± 0.1d 19 ± 2b 15 ± 2d 13 ± 1e 2.3 ± 0.0h
D. bulbifera 54 ± 1e 3 ± 0.1cd 0.58 ± 0.04d 1.22 ± 0.04c 8.6 ± 0.2d 1.4 ± 0.0e 15.4 ± 0.3c 23 ± 2c 12 ± 1e 3.2 ± 0.0e
D. glabra 70 ± 0.4c 2.5 ± 0.2d 0.56 ± 0.03d 1.02 ± 0.03d 8.6 ± 0.2d 1.3 ± 0.0e 10.2 ± 0.2d 17 ± 0.1d 16 ± 1d 2.6 ± 0.0g
D. alata 64 ± 0.3d 3.1 ± 0.2c 1.00 ± 0.02c 1.4 ± 0.0bc 9 ± 0.1d 2.1 ± 0.1c 17 ± 1b 35.3 ± 0.5b 20.1 ± 1c 3.7 ± 0.1d

Mean 70 3.5 1.0 1.4 8.7 1.8 17 27 18.1 3.5

LSD (p < 0.05) 3.7 0.7 0.2 0.2 0.4 0.09 2.1 1.9 1.5 0.12
CV (%) 2.3 9.0 7.9 4.5 1.3 0.02 5.4 5.0 3.6 1.4

Table 2
Nutritional compositions of different Dioscorea species. Data are the mean of three replications ± SD. Means followed by a common letter in the same column are not
significantly different at the 5% level by Fisher's least significance difference (LSD) test. CV coefficient of variance; dw: dry weight.
Species Sugar (mg g−1dw) Starch (mg g−1 Amylose (mg g−1 Na (mg 100 g −1
K (g 100 g −1
dw) Vitamin-C (mg100 g-1 Vitamin-E (mg100 g-1 dw)
dw) dw) dw) dw)

D. oppositifolia 53 ± 0.2a 4.3 ± 0.0a 10 ± 0.2b 60 ± 2d 1.25 ± 0.01b 5.7 ± 0.0b 0.6 ± 0.1ab
D. hamiltonii 54 ± 1a 4.4 ± 0.1a 16 ± 1a 89 ± 1a 1.03 ± 0.01e 5.7 ± 0.2b 0.7 ± 0.0a
D. pubera 53 ± 1a 4.3 ± 0.0a 16 ± 1a 67 ± 2c 1.14 ± 0.01c 9.4 ± 0.2a 0.6 ± 0.0a
D. wallichii 52.1 ± 0.5b 4.1 ± 0.0b 9.2 ± 0.5b 68 ± 2c 1.20 ± 0.01d 5.4 ± 0.3b 0.4 ± 0.0c
D. hispida 46 ± 1c 3.1 ± 0.0e 5.2 ± 0.3d 31 ± 2e 0.83 ± 0g 1.7 ± 0.3d 0.3 ± 0.0d
D. pentaphylla 35 ± 1e 3.7 ± 0.1d 6.5 ± 0.3c 79 ± 2b 1.40 ± 0.01a 4.2 ± 0.3d 0.4 ± 0.0c
D. bulbifera 38 ± 1d 4.0 ± 0.0c 7.4 ± 0.3c 66 ± 2c 1.25 ± 0.01b 4.3 ± 0.3d 0.3 ± 0.0cd
D. glabra 35 ± 0.3e 3.1 ± 0.1d 6.4 ± 0.5c 33 ± 2e 0.83 ± 0.01g 4.1 ± 0.1d 0.3 ± 0.1c
D. alata 53 ± 0.1b 4.2 ± 0.0b 8.6 ± 0.6b 55 ± 1d 1.0 ± 0.0f 5.0 ± 0.2c 0.4 ± 0.0c

Mean 46.5 4.0 9.4 61.2 1.1 5.0 0.4

LSD (p < 0.05) 1.2 0.1 1.0 5.8 0.12 0.6 0.1
CV [%] 1.7 0.02 4.8 0.5 0.5 5.1 9

Foam capacity (FC) of tuber flour was determined as a percentage spectrophotometric method of Craig, Maningat, Seib, and Hoseney
using the method of Coffmann and Garciaj (1977). The yam flour (3 g) (1989). An aqueous suspension (1%) of yam flour was made with dis-
was transferred into 50 ml cylinders. The flour samples were gently tilled water and heated in a water bath at 100 °C for 30 min with oc-
leveled and the volumes noted. Distilled water (30 ml) was added to casional shaking. The mixture was filtered using Whatman no. 42 filter
each sample and the cylinder was vigorously swirled and allowed to paper and the paste clarity of each suspension was determined by
stand for 120 min while the change in foam volume in the cylinder was measuring the percent of transmittance at 650 nm against a water blank
measured every 15 min. on a spectrophotometer (BIO Spectrophotometer, Eppendorff, Ham-
burg, Germany).
FC (%) = [(Vol. after homogenization - Vol. before homogenization)/
(Vol. before homogenization)] × 100
2.6. Statistical analysis
The iodine affinity of starch (IAS) of the flours was assayed using the
method of Kawabata, Sawayama, Nagashima, Del Rosario, and The data reported in all the tables are mean values of triplicate
Nakamura (1984). The dried flours (3 g) were added and made up to determinations (n = 3). Differences between various parameters were
30 ml dispersions using distilled water. The dispersion was stirred oc- compared using one-way analysis of variance (ANOVA) using CROPS-
casionally within the first 30 min and then filtered through Whatman TAT ver. 7.2.2007.3 (International Rice Research Institute, Manila,
no. 42 filter paper (HiMedia). A 10 ml aliquot of the filtrate was Philippines) software. The statistical significance (p < 0.05) of the
transferred into a conical flask and phenolphthalein (4 drops) added parameter means were determined using Fisher's least significance
and the filtrate was titrated with 0.1 N I2 solution until a bluish black difference (LSD) test. The standard deviations (SD) and multiple cor-
colour formed. The starch cell damage (free starch content) was cal- relation analysis were done using Microsoft Excel ver. 2007 (MS Office,
culated using the titration value and expressed as the IAS (ppm): https://products.office.com/en-us/previous-versions/microsoft-excel-
2007). Although a p < 0.05 was generally used, the authors have also
IAS (ppm) = [(VD/VA) × (VT/Ms) × (NA/100)] × 106 chosen to use 0.01 for some of the data to indicate the greater sig-
Where VD is the total volume of the dispersion, VA is the volume of the nificance of the differences. The Bray-Curtis similarity index among
aliquot used for titration, VT is the titer value, Ms is the mass of flour different Dioscorea species were obtained using a dendogram based on
used and NA is the normality of the iodine solution used. different parameters using PAST-3 (Palaeontological Statistics,
Paste clarity (PC) of flour was determined using the University of Oslo, https://folk.uio.no/ohammer/past/) software.
Principal component analysis (PCA) of different parameters was also

4
B. Padhan, et al. Food Bioscience 34 (2020) 100527

Table 3
Anti-nutritional compositions of different Dioscorea species. Data are the mean of three replications ± SD. Means followed by a common letter in the same column
are not significantly different at the 5% level by Fisher's least significance difference (LSD) test. CV: coefficient of variance; dw: dry weight.
Species Diosgenin (mg Saponin (mg g−1 α-Amylase inhibitor (AIU/ Trypsin inhibitor Phenol (mg g−1 Tannin (mg g−1 Alkaloid (mg 100 g−1
g−1 dw) dw) mg soluble starch) (TIU/mg protein) dw) dw) dw)

D. oppositifolia 4.2 ± 0.1b 1.7 ± 0.0bc 18 ± 1b 3.4 ± 0.3b 3.5 ± 0.3c 0.4 ± 0.1c 10 ± 0.3c
D. hamiltonii 5 ± 0.2b 1.6 ± 0.0c 13.5 ± 0.4c 3.5 ± 0.4b 3.2 ± 0.2c 0.3 ± 0.0c 11 ± 0.1b
D. pubera 7 ± 0a 2.1 ± 0.0a 13 ± 0.3c 3.5 ± 0.4b 3.3 ± 0.3c 0.5 ± 0.1b 12.1 ± 0.4b
D. wallichii 5 ± 0.2b 1.6 ± 0.0c 19.4 ± 0.1a 2.7 ± 0.3b 4.4 ± 0.3c 0.5 ± 0.0b 7.2 ± 0.4d
D. hispida 4.3 ± 0.1b 1.1 ± 0.0d 14.6 ± 0.5c 5.3 ± 0.1a 4.3 ± 0.3c 0.6 ± 0.0b 10 ± 0c
D. pentaphylla 5.4 ± 0.2b 1.7 ± 0.0b 15.1 ± 0.1c 3.1 ± 0.0b 5.6 ± 0.4b 0.6 ± 0.0b 11 ± 1b
D. bulbifera 6 ± 0.1a 2.0 ± 0.0b 17.3 ± 0.3b 3.6 ± 0.3b 9 ± 0.5a 0.7 ± 0.1a 15 ± 1a
D. glabra 5.2 ± 0.1b 1.5 ± 0.0c 21 ± 0.3a 4 ± 0.2b 6 ± 0.3b 0.6 ± 0.1b 16 ± 1a
D. alata 4 ± 0.1c 1.6 ± 0.0c 11.4 ± 0.4d 2.2 ± 0.1c 3.6 ± 0.1c 0.4 ± 0.0c 10 ± 0.1c

Mean 5.0 1.7 16 3.4 4.7 0.5 11.4

LSD (p < 0.05) 1.5 0.1 3 1.1 0.7 0.1 1.6

CV [%] 2.8 0.04 2.4 8.5 5.9 11.3 7.2

used to determine the sources of variability at the level of the Dioscorea among yam species and significant (p < 0.05) varietal differences were
species using XLSTAT ver. 2014.5.03 (Paris, France), which run within observed (Table 1). The wild species such as D. hamiltonii, D. pubera and
the Excel program. D. oppositifolia showed significantly higher values of WAC. The results
imply that these wild Dioscorea species are capable of holding more
3. Results and discussion water in the flours and suggest the extent of starch gelatinization (Sila &
Malleshi, 2011). The higher WAC of the tuber may prove to be useful in
3.1. Proximate composition and physico-functional parameters enhancing the applicability of the tuber flours for making various food
product where good viscosity is required (Kaur et al., 2011). The results
Proximate analysis involves the determination of the major com- were consistent with the previous reports of Sanful and Engmann
ponents of food as moisture, ash, crude fat, crude protein and crude (2016) and also from other tuber flours such as Colocasia esculenta
fiber (Gemede, Ratta, Haki, Woldegiorgis, & Beyene, 2015). Proximate (Kaur et al., 2011) and potato (Kaur et al., 2011). Similarly, FC, PC, WSI
compositions were compared among 8 wild and one cultivated Dios- and IAS were significantly (p < 0.05) different among these yam tubers
corea species from Koraput, India (Table 1). Significant differences ex- (Table 1). The range of FC, PC, WSI and IAS in wild Dioscorea species,
isted between the moisture and ash content of the fresh tubers from the such as D. hamiltonii, D. pubera and D. oppositifolia were significantly
different yam species. The value of moisture and ash content ranged (p < 0.05) higher than that of cultivated species (D. alata). However,
from 54.0 to 82.5 and 2.5–5.4% of fresh weight, respectively (Table 1). these results were also consistent with the other tuber flours such as
The moisture and ash content was significantly (p < 0.05) higher in Colocasia esculenta (Kaur et al., 2011) and potato (Kaur et al., 2011).
some wild Dioscorea species compared to the cultivated species (D. The pasting characteristics have an important role in the selection of a
alata) except D. wallichii and D. bulbifera. The moisture and ash contents suitable species for use in the industry as a thickener and binder (Kaur
of these wild yam species were lower than the earlier reported values et al., 2011). Based on these results yam tuber flours, such as D. ha-
for tropical yam species (Aruah, Uguru, & Oyiga, 2012; Bekele & miltonii, D. pubera and D. oppositifolia have a good potential to be used
Bekele, 2018; Bhandari et al., 2003). Species with low moisture content as a food ingredient in the food industry based on the higher value of
would be suitable for prolonged tuber storage and more efficient for FC, PC, WSI and IAS.
industrial processing.
There was significant (p < 0.05) differences of crude fiber and crude 3.2. Nutritional compositions
fat among the wild and cultivated Dioscorea species. The crude fibre and
fat contents were higher than reported values for other tropical yam The nutritional compositions of different yam (Dioscorea) tubers are
species (Arinathan, Mohan, & Maruthupandian, 2009; Bhandari et al., shown in Table 2. The soluble sugar, starch and amylose content of
2003; Shajeela et al., 2011; Shanthakumari et al., 2008). Some wild different Dioscorea species were significantly (p < 0.05) different. These
yam species, such as D. oppositifolia, D. hamiltonii and D. pubera showed values of sugar, starch and amylose were consistent with the values
significantly higher values of ash, crude fiber, fat and crude protein in reported for different cultivated yam tubers of Nepal and Srilanka
comparison to cultivated species (D. alata), which suggested that they (Bhandari et al., 2003; Wanasundera & Ravindran, 1994). There were
would provide essential useful minerals. Such variation among the yam significant differences between wild and cultivated Dioscorea species
species might be related to their genetic origin and geographical with respect to carbohydrate content and the range of sugar, starch and
sources where they are grown. The higher crude protein of these yam amylose in wild Dioscorea species, such as D. hamiltonii, D. pubera and D.
tubers suggested its nutritional superiority over the other wild and oppositifolia were significantly (p < 0.05) higher than that of cultivated
cultivated yam species. Crude protein content of these yam species were species (D. alata). The high carbohydrate values of these wild Dioscorea
higher than the reported species of Sri Lanka (10.16%) (Senanayake, species make them reliable for food security crops. Significant
Ranaweera, Bamunuarachchi, & Gunaratne, 2012); Ethiopia (9.7%) (p < 0.05) variations of selected vitamin compositions was observed
(Tamiru, Mass, & Pawelzik, 2008); Ghana (4.0–6.5%) (Polycarp et al., among the studied Dioscorea species. The low value of vitamin C con-
2012), Indonesia (5.3%) (Aprianita, Vasiljevic, Bannikova, & Kasapis, tent was believed to be due to the heat destruction with yam processing.
2014), cassava (1.2–1.8%) (Charles, Sriroth, & Huang, 2005) and sweet Based on these results, some wild yam species, such as D. hamiltonii, D.
potato (5.6%) (Moongngarm, 2013) but lower than some Indian yam pubera and D. oppositifolia showed significantly (p < 0.05) higher vi-
species (15.8%) (Shanthakumari et al., 2008). tamin content and suggested its nutritional superiority over the other
The physico-functional properties such as WAC, FC, PC, WSI and Dioscorea species. However, the yam tubers showed higher vitamin C
IAS are the important parameters for bioprospecting of the tuber flour content than the earlier report in the tubers of D. alata (Mohan &
(Kaur, Kaushal, & Sandhu, 2011). These parameters were compared Kalidass, 2010; Udensi, Oselebe, & Iweala, 2008). There were

5
B. Padhan, et al. Food Bioscience 34 (2020) 100527

Multiple correlation between physico-functional parameters with nutritional and phenol content in different Dioscorea species. Phe: phenol; Sug: sugar; Sta: starch; Amy: amylose; Pro: protein; Vit: vitamin; Cfat: Crude fat;
Cfib: Crude fibre Mc: moisture content; WAC: water absorption capacity; FC: foam capacity; PC: paste clarity; WSI: water solubility index; IAS: iodine affinity of starch; Dio: Diosgenin. Degree of freedom (df) = 26; *: significant (p < 0.05) differences of mineral compositions such as Na

ns
0.35
and K content among different Dioscorea species (Table 2). The level of
K
K was higher than the Na content in the studied yam tubers, which have
better health implication and considered as a safe food for human

0.29 ns
0.70**
consumption (Appiah, Asibuo, & Kumah, 2011). The proper balance of
Na

K in the body depends on Na intake and the daily intake of K between

1875 and 5625 mg is considered to be adequate and safe, however, the

−0.34 ns
0.32 ns
0.13 ns
K content of these yam tubers was less than the critical limits (Baah,
PC

Maziya-Dixon, Asiedu, Oduro, & Ellis, 2009) and is recognising that

these would not be the sole, but a major source of K in the diet of these

0.24 ns
0.52**
0.68**
0.65**
native people.
FC

3.3. Anti-nutritional compositions

0.30 ns
0.17 ns
0.25 ns
0.62**
0.63**
IAS

Anti-nutritional compositions are the chemical compounds synthe-

sized in natural food by normal metabolism which decreases the nu-
tritive value of food (Ugwu & Oranye, 2006; Fekadu, Beyene, & Desse,

−0.05 ns
0.26 ns
0.05 ns
0.87**
0.56**
0.72**

2013). The anti-nutritional factors, such as diosgenin, phenols, tannins,

WSI

saponin and alkaloids contents of different Dioscorea species are shown

in Table 3. Diosgenin is one of the bioactive compounds used in the
ns

synthesis of steroidal drugs and is the primary active ingredient of

0.70**
0.57**
0.84**
0.74**
0.72**

−0.09
0.41*
WAC

Dioscorea species (Adepoju et al., 2018). The diosgenin content was

significantly (p < 0.05) higher in different wild Dioscorea species
compared to the cultivated species (D. alata). The results were con-
−0.02 ns

sistent with the earlier reported values for different tropical yam species
0.15 ns
0.79**
0.72**
0.66**
0.57**
0.70**
0.60**
Ash

(Adepoju et al., 2018). Similarly, phenol, tannin, saponin and alkaloid

content were significantly (p < 0.05) higher in some wild Dioscorea
species compared to the cultivated one (Table 3). Further, the α-amy-
−0.06 ns
−0.25 ns

−0.35 ns

−0.16 ns
−0.01 ns
0.06 ns
0.21 ns

0.16 ns

0.07 ns

lase and trypsin inhibitor were significantly (p < 0.05) varied among
MC

the wild and cultivated Dioscorea species (Table 3). The range of these
parameters at low levels may have a beneficial role as an antioxidant,
−0.24 ns

anticarcinogens and likely have an important role in controlling hy-

0.07 ns
0.55**
0.54**
0.70**
0.72**
0.53**

0.54**
0.43*

0.45*

percholesterolemia and atherosclerosis (Phillippy, Lin, & Rasco, 2004).

Cfib

The values of different anti-nutritional compositions in the yams of

Koraput were lower than the value reported by Polycarp et al. (2012) in
0.12 ns

0.15 ns
0.01 ns
0.94**
0.85**
0.79**
0.69**
0.64**
0.74**
0.53**
0.46*

Ghanaian Dioscorea species. Such variations of anti-nutritional compo-

Cfat

sitions might be related to their genetic origin, geographical sources,

level of soil fertility and the harvesting periods (Polycarp et al., 2012).
0.24 ns

0.26 ns
0.14 ns
0.93**

0.90**
0.86**
0.72**
0.61**
0.68**
0.61**
0.66**
0.48*
Vit E

3.4. Relationship between physico-functional parameters with nutritional

and anti-nutritional compositions
ns
0.61**
0.59**
0.57**

0.51**
0.57**
0.70**
0.84**
0.70**

0.57**
−0.22

0.47*
0.47*
0.41*
Vit C

Relationship between various physico-functional parameters with

nutritional and antinutritional parameters was studied using multiple
correlation analysis (Table 4). The results showed that physico-func-
−0.03 ns
0.01 ns

0.01 ns
0.58**
0.82**
0.89**
0.69**

0.90**
0.74**
0.87**
0.83**
0.53**
0.70**
0.46**

tional properties such as IAS, WSI and PC were not significantly

Pro

(p ≥ 0.05) influenced by mineral composition (Na and K) and moisture

content but there was a significant (p < 0.01) positive association with
0.11 ns

0.14 ns
0.86**
0.78**
0.88**
0.87**
0.51**

0.83**
0.75**
0.75**
0.78**
0.65**
0.56**
0.60**

WAC and PC of yam tuber. The physico-functional parameters were

0.37*
Amy

significantly positively correlated with sugar, starch, amylose, crude

protein, vitamins, crude fat, and fiber, whereas phenol content was
ns

ns

negatively correlated. The high phenolic compound could affect the FC

0.63**
0.69**
0.67**
0.64**
0.66**
0.76**

0.68**
0.81**
0.67**
0.71**
0.76**
0.81**
0.70**
0.55**
−0.35

−0.02

and PC by destabilising the protein films surrounding the air droplets

p < 0.05, **: p < 0.01, ns: non-significant.
Sta

and causing the foam to collapse as reported by Craig et al. (1989).

−0.12 ns

−0.34 ns

3.5. Principal component analysis and cluster analysis

0.33 ns
0.13 ns
0.77**
0.53**
0.78**

0.50**
0.64**
0.64**

0.65**
0.70**
0.69**
0.72**
0.59**
0.76**
0.38*
Sug

The data for the different parameters were subjected to multivariate

analysis including PCA and cluster analysis. The first two axes of the
−0.33 ns

−0.29 ns
−0.27 ns

−0.09 ns
−0.65**

−0.62**

−0.49**
−0.58**

−0.55**
−0.62**
−0.62**

−0.53**
−0.33ns
−0.33ns

−0.39*

0.26 ns
0.67**

principal component analysis captured 70.6% of the total variation

0.03ns
Phe

with eigen values more than one, suggest the wide genetic variation
among the studied Dioscorea species (Fig. 2). The PC1 accounted for the
Parameters

majority of the variation (57.3%) of studied parameters. In PC1, mi-

nerals and vitamin compositions are the dominant variables showed the
Table 4

WAC
Vit E
Amy

VitC

Cfib
Cfat

WSI
Ash

highest positive value, whereas in PC2 was constituted mainly positive

Sug

Dio
IAS
Pro

MC
Sta

Na
PC
FC

effects of anti-nutritional compositions, such as phenol, tannin and

6
B. Padhan, et al. Food Bioscience 34 (2020) 100527

they showed similar tuber quality traits.

4. Conclusion

The tuber quality traits between wild and cultivated Dioscorea

species showed that some wild species had better nutritional compo-
sition and mineral content in comparison to the cultivated species.
Some of the wild Dioscorea species were found to be a good source of
vital nutrients such as carbohydrate, protein, crude fiber and K. The
wild species, such as D. hamiltonii, D. pubera and D. oppositifolia had
significantly higher amounts of nutritional and mineral content and
suggesting it was nutritionally better than other. The physico-functional
parameter of this tuber flour was also better. The findings of the study
also suggested that these less familiar wild tubers should not be ig-
nored. Rather, these tubers are safe food sources with a good potential
for domestication.
Fig. 2. Biplot of principal component (PC) analysis for different Dioscorea
species based on nutritional, anti-nutritional and physico-functional para- CRediT authorship contribution statement
meters.

Bandana Padhan: Formal analysis, Writing - original draft.

Meghali Biswas: Formal analysis, Writing - original draft. Debabrata
Panda: Formal analysis, Writing - original draft.

Declaration of competing interest

All authors have declared that they do not have any conflict of in-
terest for publishing this research.

Acknowledgements

Authors are grateful to the Head, Department of Biodiversity and

Conservation of Natural Resources for providing necessary facilities for
the study and also grateful to the University Grant Commission (UGC),
New Delhi, India for providing a Non-NET Fellowship.

Appendix A. Supplementary data

Supplementary data to this article can be found online at https://

doi.org/10.1016/j.fbio.2020.100527.

Fig. 3. Dendrogram showing the percentage of similarity between wild and References
cultivated species of Dioscorea based on nutritional, anti-nutritional and phy-
sico-functional parameters. Adebooye, O. C., & Phillips, O. T. (2006). Studies on seed characteristics and chemical
composition of three morphotypes of Mucuna urens (L.) Medikus. Fabaceae. Food
Chemistry, 95(4), 658–663.
alkaloid. Based on the loading and biplot, it was seen that a few traits Adepoju, O. T., Boyejo, O., & Adeniji, P. O. (2018). Effects of processing methods on
are the major determinants of phenotypic diversity, among which nutrient and antinutrient composition of yellow yam (Dioscorea cayenensis) products.
Food Chemistry, 11, 428–431.
physico-functional parameters such as WAC, WSI and PC have an im- Alonso, R., Orúe, E., & Marzo, F. (1998). Effects of extrusion and conventional processing
portant role. The rest of the compositions seem to have a very minimal methods on protein and antinutritional factor contents in pea seeds. Food Chemistry,
contribution to variability. A scatter plot was drawn between PC1 and 63(4), 505–512.
Amanze, N. J., Agbo, N. J., Eke-Okoro, O. N., & Njoku, D. N. (2011). Selection of yam
PC2 that showed the pattern of grouping different Dioscorea species in
seeds from open pollination for adoption in yam (Dioscorea rotundata Poir) produc-
the factor plane. Based on these results, wild yam species, such as D. tion zones in Nigeria. Journal of Plant Breeding and Crop Science, 3(4), 68–73.
hamiltonii, D. pubera, D. oppositifolia and D. wallichii along with culti- Anderson, R. A., Conway, H. F., Pfeifer, V. F., & Griffin, E. L. (1969). Roll and extrusion-
vated species D. alata present in one quarter and were loaded to the cooking of grain sorghum grits. Cereal Science Today, 14(11), 372–380.
AOAC (2012). Official methods of analysis of the association of official analytical chemists
positive side of PC1, where as D. hispida and D. glabra were in the (19th ed.). Gaithersburg, MD, USA: AOAC International.
second quarter and loaded to the negative end of PC1 (Fig. 2). D. bul- Appiah, F., Asibuo, J. Y., & Kumah, P. (2011). Physicochemical and functional properties
bifera and D. pentaphylla had a negative score in the second principal of bean flours of three cowpea (Vigna unguiculata L. Walp) varieties in Ghana. African
Journal of Food Science, 5(2), 100–104.
component (PC2). Cluster analysis based on the Bray-Curtis paired Aprianita, A., Vasiljevic, T., Bannikova, A., & Kasapis, S. (2014). Physicochemical prop-
linkage showed the percent of similarity in tuber quality parameters erties of flours and starches derived from traditional Indonesian tubers and roots.
among wild and cultivated Dioscorea species (Fig. 3). The dendrogram Journal of Food Science & Technology, 51, 3669–3679.
Arinathan, V., Mohan, V. R., & Maruthupandian, A. (2009). Nutritional and antinutri-
showing the similarity forming two major clusters. Cluster I was further tional attributes of some under –utilized tubers. Tropical and Subtropical
divided into two subgroups which included wild yam species, such as D. Agroecosystem, 10, 273–278.
hamiltonii, D. pubera D. oppositifolia and D. wallichii along with the Aruah, B. C., Uguru, M. I., & Oyiga, B. C. (2012). Genetic variability and inter-relationship
among some Nigerian pumpkin accessions (Cucurbita spp). International Journal of
cultivated species D. alata in one cluster having more than 92% simi- Plant Breeding, 6(1), 34–41.
larity, whereas, D. pentaphylla and D. bulbifera were in a separate sub- Baah, F. D., Maziya-Dixon, B., Asiedu, R., Oduro, I., & Ellis, W. O. (2009). Nutritional and
cluster. The species which are within the same cluster are unique as biochemical composition of D. alata (Dioscorea spp.) tubers. Journal of Food
Agriculture and Environment, 7(2), 373–378.

7
B. Padhan, et al. Food Bioscience 34 (2020) 100527

Baker, H., Frank, O., Angelis, B. D., & Feingold, S. E. (1980). Plasma tocopherol in man at USA: Academic Press.
various times after ingesting free or acetylated tocopherol. Nutrition Reports Otegbayo, B. O., Oguniyan, D. J., Olunlade, B. A., Oroniran, O. O., & Atobatele, O. E.
International, 21(4), 531–536. (2018). Characterizing genotypic variation in biochemical composition, anti-nutri-
Behera, K. K., Maharana, T., Sahoo, S., & Prusti, A. (2009). Biochemical quantification of tional and mineral bioavailability of some Nigerian yam (Dioscorea spp.) land races.
protein, fat, starch, crude fibre, ash and dry matter content in different collection of Journal of Food Science & Technology, 55(1), 205–216.
greater yam (Dioscorea alata L.) found in Orissa. Nature and Science, 7(7), 24–32. Padhan, B., & Panda, D. (2016). Wild tuber species diversity and its ethno-medicinal use
Bekele, A., & Bekele, E. (2018). Proximate and mineral composition variability in by tribal people of Koraput district of Odisha, India. Journal of Natural Products and
Ethiopian yam (Dioscorea spp). Journal of Food and Nutrition Sciences, 6(1), 12–17. Resources, 2(1), 33–36.
Bhandari, M. R., Kasai, T., & Kawabata, J. (2003). Nutritional evaluation of wild yam Phillippy, B. Q., Lin, M., & Rasco, B. (2004). Analysis of phytate in raw and cooked
(Dioscorea spp.) tubers of Nepal. Food Chemistry, 82, 619–623. potatoes. Journal of Food Composition and Analysis, 17(2), 217–226.
Bhandari, M. R., & Kawabata, J. (2004). Assessment of antinutritional factors and bioa- Phillips, R. D., Chinnan, M. S., Branch, A. L., Miller, J., & Mcwatters, K. H. (1988). Effects
vailability of calcium and zinc in wild yam (Dioscorea spp.) tubers of Nepal. Food of pre-treatment on functional and nutritional properties of cowpea meal. Journal of
Chemistry, 85, 281–287. Food Science, 53(3), 805–809.
Charles, A. L., Sriroth, K., & Huang, T. C. (2005). Proximate composition, mineral con- Polycarp, D., Afoakwa, E. O., Budu, A. S., & Otoo, E. (2012). Characterization of chemical
tents, hydrogen cyanide and phytic acid of five cassava genotypes. Food Chemistry, composition and anti-nutritional factors in seven species within the Ghanaian yam
92, 615–620. (Dioscorea) germplasm. International Food Research Journal, 19(3), 985–992.
Coffmann, C. W., & Garciaj, V. V. (1977). Functional properties and amino acid content of Price, E. J., Bhattacharjee, R., Montes, A. L., & Fraser, P. D. (2017). Metabolite profiling of
a protein isolate from mung bean flour. Journal of Food Technology, 12(5), 473–484. yam (Dioscorea spp.) accessions for use in crop improvement programmes.
Craig, S. A. S., Maningat, C. C., Seib, P. A., & Hoseney, R. C. (1989). Starch paste clarity. Metabolomics, 13(144), 1–12.
Cereal Chemistry, 66(3), 173–182. Sadasivam, S., & Manickam, A. (2007). Biochemical methods. New Delhi, India: New Age
Fekadu, H., Beyene, F., & Desse, G. (2013). Effect of traditional processing methods on International Pvt. Ltd284.
nutritional composition and anti-nutritional factors of anchote (Coccinia Abyssinica Sanful, R. E., & Engmann, F. N. (2016). Physico-chemical and pasting characteristics of
(lam.) Cogn) tubers grown in Western Ethiopia. Journal of Food Processing & flour and starch from aerial yam. American. Journal of Food Science and Nutrition,
Technology, 4(7), 249. 3(1), 1–7.
Gemede, H. F., Ratta, N., Haki, G. D., Woldegiorgis, A. Z., & Beyene, F. (2015). Nutritional Senanayake, S. A., Ranaweera, K. K. D. S., Bamunuarachchi, A., & Gunaratne, A. (2012).
quality and health benefits of okra (Abelmoschus esculentus): A review. Journal of Food Proximate analysis phytochemical and mineral constituents in four cultivars of yams
Processing & Technology, 6(6), 458. and tuber crops in Srilanka. Tropical Agricultural Research and Extension, 15, 32–36.
Harborne, J. B. (1973). Phytochemical methods: A guide to modern techniques of plant Shajeela, P. S., Mohan, V. R., Jesudas, L. L., & Soris, P. T. (2011). Nutritional and anti-
analysisLondon: Chapman and Hall Ltd1–286. nutritional evaluation of wild yam (Dioscorea spp.). Tropical and Subtropical
Kaur, M., Kaushal, P., & Sandhu, K. S. (2011). Studies on physicochemical and pasting Agroecosystems, 14, 723–730.
properties of taro (Colocasia esculenta L.) flour in comparison with a cereal, tuber and Shanthakumari, S., Mohan, V. R., & Britto, J. De (2008). Nutritional evaluation and
legume flour. Journal of Food Processing & Technology, 50(1), 94–100. elimination of toxic principle in wild yam (Dioscorea spp.). Tropical and Subtropical
Kawabata, A., Sawayama, S., Nagashima, N., Del Rosario, R. R., & Nakamura, M. (1984). Agroecosystem, 8, 313–319.
Some physico-chemical properties of starches from cassava, arrow root and sago. Sila, B., & Malleshi, N. G. (2011). Physical, chemical and nutritional characteristics of
Journal of Japanese Society of Starch Science, 31, 224–232. premature-processed and matured green legumes. Journal of Food Science &
Kouakou, M. D., Dabonne, S., Guehi, T., & Kuoame, L. P. (2010). Effects of post-harvest Technology, 49(4), 459–466.
storage on some biochemical parameters of different parts of two yams species Smith, C., Megen, W. V., Twaalfhoven, L., & Hitchcock, C. (1980). The determination of
(Dioscorea spp). African Journal Food Science and Technology, 1, 1–9. trypsin inhibitor levels in foodstuffs. Journal of the Science of Food and Agriculture, 31,
Mishra, S., & Chaudhury, S. S. (2012). Ethnobotanical flora used by four major tribes of 341–350.
Koraput, Odisha, India. Genetic Resources and Crop Evolution, 59(5), 793–804. Tamiru, M., Mass, B. L., & Pawelzik, E. (2008). Characterizing diversity in composition
Mishra, S., Swain, S., Chaudhary, S., & Ray, S. (2011). Wild edible tubers (Dioscorea spp.) and pasting properties of tuber flour in yam germplasm (Dioscorea spp.) from
and their contribution to the food security of tribes of Jeypore tract, Orissa, India. Southern Ethiopia. Journal of the Science of Food and Agriculture, 88, 1675–1685.
Plant Genetic Resources Newsletter, 56, 63–67. Udensi, E. A., Oselebe, H. O., & Iweala, O. O. (2008). The investigation of chemical
Mohan, V. R., & Kalidass, C. (2010). Nutritional and antinutritional evaluation of some composition and functional properties of water yam (Dioscorea alata): Effect of var-
unconventional wild edible plants. Tropical and Subtropical Agroecosystems, 12, ietal differences. Pakistan Journal of Nutrition, 7(2), 342–344.
495–506. Uematsu, Y., Hirata, K., Saito, K., & Kudo, I. (2000). Spectrophotometric determination of
Moongngarm, A. (2013). Chemical compositions and resistant starch content in starchy saponin in Yucca extract used as food additive. Journal of AOAC International, 83(6),
foods. American Journal of Agricultural and Biological Sciences, 8, 107–113. 1451–1454.
Nahapetian, A., & Bassiri, A. (1975). Changes in concentration and interrelationship of Ugwu, F. M., & Oranye, N. A. (2006). Effects of some processing methods on the toxic
phytate, P, Mg, Cu, Zn, in wheat during maturation. Journal of Agricultural and Food components of African bread fruit (Treculia africana). African Journal of Biotechnology,
Chemistry, 32, 1179–1182. 5, 2329–2333.
Ofosu, S. B. (2012). Assessment of three white yam (Dioscorea rotundata) varieties for possible Wanasundera, J. P. D., & Ravindran, G. (1994). Nutritional assessment of yam (Dioscorea
development into flourM.Phil. Thesis. Kumasi, Ghana: Kwame Nkrumah University of alata) tubers. Plant Foods for Human Nutrition, 46(1), 33–39.
Science and Technology1–4. Wu, Z. G., Wu, J., Nitin, M., Bao, X. Q., Chen, S. L., & Tao, Z. M. (2016). Characterizing
Omaye, S. T., Turnbull, T. D., & Sauberlich, H. E. (1979). Selected method for the de- diversity based on nutritional and bioactive compositions of yam germplasm
termination of ascorbic acid in animal cells, tissues and fluid. In D. B. Mc Cormic, & (Dioscorea spp.) commonly cultivated in China. Journal of Food and Drug Analysis, 24,
D. L. Wright (Vol. Eds.), Methods in enzymology: Vol. 62, (pp. 3–11). New York, NY, 367–375.