The Thirty-nine genotypes of Coix showed high variability in terms of yield and yield related traits. Characterization and evaluation of germplasm are crucial for producing improved varieties (Nelson 2011; Andini et al. 2013). Thus, evaluation of germplasm to identify potential genotypes for cultivation was done for eight traits. The descriptive statistics have been demonstrated in our present study for eight traits and are presented in Table 1.
Descriptive statistics
Significant variations were observed among genotypes for different traits. The highest coefficient of variation was reported in plant height (22.10%) followed by LAI (8.38%) and seed yield (5.05%). The plant height among the genotypes varies from 118.63 cm (RJTGP-51) to 237.33 cm (RJTGP-59), with a mean value of 174.54 cm. A key factor in determining the appropriate shoot character is plant height. In many cereal crops, like rice, wheat and maize, shorter stem contributes to higher yield through improved resistance to lodging (Chairi et al. 2020; Chen et al. 2018; Su 2018; Cooper et al. 2003; Cooper et al. 1995). At present, the optimum height of Coix for ideal shoot architecture is yet to be exploited to develop high yielding varieties. The number of leaves/plants varies from 24.96–100.00, with a mean value of 49.15 leaves/plant. Number of leaves is an important parameter to be considered for fodder purpose. RJTGP-90 has the maximum number of leaves followed by RJTGP-89 (87.87) and RJTGP-71 (80.00). Leaf area was ranging from 685.92-2742.35 cm2 with a mean value of 1754.89 cm2. RJTGP-90 exhibited maximum leaf area while IC-540173 exhibited minimum leaf area. The LAI was ranging from 1.19–4.72, with a mean value of 3.03. LAI helps in understanding photosynthesis, light interception, nutrient and water utilization, growth of crop and potential of yield (Smart 1974; Williams 1987). The highest LAI was exhibited by RJTGP-90 followed by RJTGP-57 (4.61). The number of tillers/plant varies from 2.08 (RJT-2) to 6.20 (IC-540173), with a mean value of 3.91. The number of seeds/plant varies from 22.22–245.00. The maximum number of seeds/plant was reported in IC-540256 (245.00) followed by RJTGP-71 (183.00) and RJTGP-74 (150.00) while the minimum was reported in IC-12703 (22.22). The maximum 100 seed weight was reported in RJTGP-71 (34.60 g) followed by RJTGP-53 (32.84 g) and IC-417053 (30.91 g), while the minimum was reported in IC-540279 (12.96 g) with a mean value of 23.59 g among the genotypes. As seed is the economical part of the crop, the selection of genotypes with high seed yield will be beneficial. The variation in seed yield ranges from 3.28–58.43 q/ha with a mean value of 16.64 q/ha. Four genotypes viz., RJTGP-53 (58.43 q/ha), RJTGP-71 (39.96 q/ha), RJT-1 (39.74 q/ha) and RJT-2 (35.27 q/ha) were found to contribute maximum for seed yield while the minimum seed yield was reported in IC-89384 (3.28 q/ha). Hore and Rathi (2006) characterized 29 genotypes of Coix collected from Northeast India and found great variability with respect to yield and nutritional content and suggested genotypes IC-012703 (31.8 q/ha), IC-089391 (30 q/ha) and IC-521338 (29 q/ha) were promising for cultivation.
Principal component analysis (PCA)
The purpose of this analysis was to determine the factor dimension of the data, which was used to simplify varietal information into a smaller number of factors and selecting the genotype(s) that would perform best for eight yield and yield-related traits. With eigen values greater than one, the first three principal components (PCs) explained 75.78% of the total variance, with PC1 accounting for 33.60% and PC2 accounting for 24.54% (Fig. 4). From the biplot figure, traits like PH, NLPP, LA and LAI were found to have high positive association with PC1 whereas SY, HSW and NSPP exhibit positive association with PC2. NTPP was positioned close to the center of the biplot, which showed relative similar performance by the genotypes. Among those genotypes, RJTGP-90, RJTGP-50, RJTGP-89, RJTGP-92 and IC-540279 were identified as superior genotypes in PC1. In contrast, RJTGP-53, RJTGP-71, RJT-1, RJT-2, RJTGP-51, and RJTGP-81 were identified as superior genotypes in the PC2.
Correlation analysis
The correlation analysis determines the degree of character association between the various traits and sgenerate information that are needed to be considered in selection programme as selection of one or more traits affects the expression of several other traits. Pearson correlation coefficients among eight traits in the current study were shown in Fig. 5. The highest correlation coefficient was observed between LA and LAI (0.99**), followed by NSPP and SY (0.73**) and LAI and NLPP (0.56**). For yield traits, NSPP and HSW showed highly significant correlation with SY while traits like PH, LA, LAI and NTPP showed negative correlation with SY, however, exhibited insignificant values. Selection of genotypes which possess high NSPP and HSW will be beneficial for improving yield of crop.
Stability analysis
The multi-trait stability index (Olivoto et al. 2021) has been successfully used to select genotypes for multiple traits based on mean performance and stability. To choose the stable genotype, most plant breeders used traditional stability criteria such the mean, regression, and deviation from regression parameters (Abdelmohsen 2021). However, these statistical tools are inconsistent to identify strengths and weaknesses of a genotype and aid in selection of those with desired mean performance and stability (Olivoto et al. 2021). Based on MTSI analysis, out of forty-four genotypes, six genotypes, RJTGP-71, RJT-2, RJT-1, RJTGP-90, RJTGP-81 and RJTGP-50 were found most stable with desired mean performance under Mizoram condition for representing a selection intensity of 10% for yield and yield related traits. These genotypes cross the cut-off point (red circle) as presented in Fig. 6(a).
The MGIDI index representing strengths and weaknesses view of the selected genotypes was given in Fig. 6(b) & Table 2. Each factor contribution to the MGIDI index is ranked from the most contributing factor (near to plot center) to the less contributing factor (near to the plot edge). The first factor FA1 (NLPP, LA, LAI) has smallest contribution for genotypes RJTGP-90 and RJTGP-57 indicating that these genotypes presented higher values in terms of number of leaves/plant, leaf area and LAI. While considering these genotypes in FA2, they had higher contribution to MGIDI which resulted from their poor performance for seed yield/plant, number of seeds/plant and 100 seed weight. On the contrary, genotypes like RJTGP-53, RJT-1 and RJT-2 performed well for these traits in FA2 (Fig. 7). The FA3 (PH, NTPP) has smallest contribution for genotypes like RJTGP-71, IC-540256 and RJTGP-50, indicating their better performances for plant height and number of tillers/plant.
Table 2
Factors linked to correlated traits, selection differential, selection differential percentage and indicators for Job’s tears traits.
Traits | Factor | Indicator | Xo | Xs | SD | SDperc |
NLPP | FA1 | increase | 49.90 | 75.70 | 25.80 | 51.60 |
LA | FA1 | increase | 1789.00 | 2161.00 | 372.00 | 20.80 |
LAI | FA1 | increase | 3.09 | 3.74 | 0.65 | 21.00 |
NSPP | FA2 | increase | 79.90 | 122.00 | 42.50 | 53.10 |
HSW | FA2 | increase | 24.20 | 28.40 | 4.26 | 17.70 |
SY | FA2 | increase | 18.20 | 28.80 | 10.60 | 58.40 |
PH | FA3 | increase | 173.00 | 173.00 | -0.43 | -0.25 |
NTPP | FA3 | increase | 3.99 | 4.11 | 0.12 | 3.00 |
Xo: original value; Xs: selected value; SD: selection differential; SDperc: selection differential in percentage. |
Based on MTSI and MGIDI analyses, these genotypes RJT-1 and RJT-2 gathered desirable overall performance and stability of several traits such as seed yield, number of seeds/plant and 100 seed weight. The higher seed yield/plant of these genotypes are the result of higher number of seeds/plant and 100 seed weight due to the highly significant positive association between the traits. It is imperative to select genotypes stable in performance with acceptable yield than genotypes with high yield and low stability for varietal recommendation. The high yielding unstable genotype like RJTGP-53 can be considered for location specific adaptation and can used in crop improvement programme with the help of stable genotypic background.
Importance Of Coix
Coix has multiple uses such as food, medicinal and traditional as follows:
Coix as an important food source
Coi x can be used in multiple forms viz., as a pseudo cereal, as a rice substitute in all dishes, and in cakes, bread, and pastries when the grain is roasted before being husked. The gluten-free nature of Coix flour makes it significant in bakery products to be used along with wheat flour (Keeratibunharn and Krasaekoopt 2013). Gluten-free products prepared from Coix grains will benefit people having celiac disease (Comino et al. 2013). It can also be prepared as soups, porridge, gruels, salads, desserts, eaten like peanuts and used for the preparation of sweets. Matured dehulled seeds can also be cooked and consumed along with rice. They can be used as snacks of different flavours by cooking, drying, deep frying and baking. It is utilized by the Japanese and Nagas to make alcoholic beverages like a beer called ‘Zhu’ or ‘Dzu’ by fermenting whole seeds (Hore and Rathi 2006). Coix seeds along with glutinous seeds can additionally be used to make fermented glutinous wine (Wu 2010). It is also consumed as a drink or as a tea by grinding the seeds and mixing or boiling with water. Vinegar can also be prepared from coix seeds. Coix is also found to be a nutritious feed for animals. Poultry is given the entire grain, bran, and flour as poultry feed in the replacement of maize. Growing in the wild, its leaves are highly preferred and consumed by wild elephants. Coix is used as feed for cattle, buffaloes and horses as a fodder crop and has better potential to be used in both summer and dry winter seasons with proper management strategies since the grass can be used to produce silage (Jain and Banerjee 1974). In addition, the seeds can improve performance in growth and productivity by lowering gut pH and regulating the gut microbiota of post-weaning pigs when fed as a feed supplement (Li et al. 2019). This crop can be harvested in a short duration, easy to cultivate and need fewer requirements as compared to other cereal crops.
Coix - an excellent source of nutrients
The grains of Coix are found to be higher in protein, fat and fibre as compared to many other cereal crops. Coix grain contains 62.0% total starch content, 15.9% protein, 4.66% crude fat, 5.53% dietary fibre, 1.52% total soluble sugar, 0.73% GAE total phenol content, 0.24% phytate, 0.47% GAE antioxidant activity, and also 0.80 mg/100 g, 146 mg/100 g, 3.61 mg/100 g, and 13.60 mg/100 g of copper, calcium, zinc and iron, respectively (Laxmisha et al. 2022). It contains 100.25, 80.68, 50.00 and 44.55% higher protein than rice, maize, wheat and pearl millet, respectively. When compared to rice, it contains a calcium level that is twelve times higher. It also has 96.80% more dietary fibre than rice. It is also an excellent source of iron as compared to rice (0.65 mg/100 g), maize (2.49 mg/100 g), wheat (3.97 mg/100 g) and pearl millet (6.42 mg/100 g) (Longvah et al. 2017). As far as the nutrient composition is concerned, we can clearly conclude that Coix is a rich source of nutrients as compared to many staple food crops of the world and the inclusion of these healthy grains in our diet can address the nutrient deficiency problem around the globe. In the Philippines, the government is promoting Coix because of its high protein content and other nutrients including calcium, phosphate, iron, thiamine, riboflavin, and niacin (Magpantay et al. 2021).
Coix for medicinal uses
Coi x plant has been used for a long time and is documented in Indian Ayurveda and traditional Chinese medicine. It consists of many miraculous medicinal properties. It is known to be useful in removing kidney and bladder stones (Hore and Rathi 2006). Furthermore, it is taken due to its effectiveness in treating respiratory tract infections, appendicitis, and allergic disorders. Its use for women as a remedy to a menstrual disorder such as excessive bleeding or irregular menstruation along with its effect in stimulating ovulation and inducing fertility is also reported. Recent research studies on the anti-cancer properties of Coix have shown encouraging results against cancers of the blood, breast, colon, liver, lung, and pancreas (Kumar et al. 2014). From the seed oil of Coix, ‘Kanglaite’ an anticancer medication, was developed. The drug is currently being investigated in the USA as a possible treatment for prostate and pancreatic cancers. It is already approved in China to treat a number of cancers (Xi et al. 2016). Coixenolide, a fatty acid ester having anticancer action, is the main bioactive component (Ukita and Tanimura 1961). Several studies have demonstrated that regular intake of grains of Coix balances human blood cholesterol levels and lowers the risk of cardiovascular diseases like atherosclerosis and myocardial infarction (Zeng et al. 2021). It is also reported to have diuretic and blood-purifying properties (Schaaffhausen, 1952). Its demand has been rising day by day as a result of its high medicinal properties as an ideal ‘functional food’.
In Mizoram, the rural communities have been using decoctions of roots as well as leaves for treating urinary tract infections. Reports show that decoction of the roots is being used to treat dysentery, gonorrhoea and as a vermifuge. The sap from the stem is utilized to treat insect bites, while a decoction of the leaves is used to treat rheumatism, headache and diabetes. The seeds of coix can revitalize the spleen, expel mucus and pus, drains dampness, clear heat, get rid of edema, and decrease body weight (Yu et al. 2011). The seeds are additionally reported to be used in treating arthritis, neuralgia, warts and chapped skin (Shih et al. 2004). The fruits have diuretic, anti-inflammatory, antidiarrheal, anti-oxidant, antipyretic, antiseptic, antispasmodic, analgesic, hypotensive, and sedative properties. Contrary to its multiple health benefits, caution is advised during pregnancy and breastfeeding as the seeds of the plant have abortifacient activity (Patel et al. 2017).
Coix as traditional items and other uses
Coi x gives the most perfect beads for various ornamental purposes. The seeds obtained from wild types have been used by rural folks to make different kinds of ornamentals –earrings, rosaries, garlands, rings, bracelets, belts, photo frames, and curtains (Fig. 7). They are used in fashion apparel by sewing the beads onto bags, jackets and head gears. The traditional head gear of the Mizo women called ‘Vakiria’ is decorated with Coix seeds. The beads of Coix can also be of value in making musical instruments such as a Shaker’s gourd by covering hollow gourds with a loose net strung with beads of coix (Biswas and Das 2022). They may be woven into baskets and vessels to add to their aesthetics. Coix seeds are an excellent source of natural jewellery as it has a hole in the centre through which wire or tread can easily pass from it. The roots are reportedly used for rituals to contain the spread of smallpox (Jain and Banerjee 1974). The stems can also be used to make mattings. A type of compost can also be made from the leaves by leaving them in piggeries, trampling and mixing with mud (Arora 1977). Additionally, its straw and leaves are utilized for thatching purposes. Moreover, this plant is a good choice for wastewater treatment in tropical areas because it is effective at eliminating inorganic nitrogen from polluted waterways (Jampeetong et al. 2013).
Cultivation Practices
Coi x is propagated by seeds and cultivated as an annual crop. It is a macrophyte that prefers soil moisture. Land selection for sowing should be low-lying with proper drainage facilities. In case of moisture stress, supplementary irrigation may be given in the early growth and flowering stage. The climatic requirement for Coix is similar to maize and can withstand wet or waterlogged conditions for a short time span. Sowing was done during monsoon preferably from May to June and sometimes in July at a seed rate of 6–10 kg/ha having a 1000-grain weight of 80–90 g. Seed dibbled 4–5 cm deep with a spacing of 60 × 60 cm or 60 × 45 cm or 45 × 15 cm. 40 kg N, 20 kg P2O5, and 20 kg K2O are recommended as general dosages of NPK per hectare. From this, 20 kg of nitrogen and a full dose (20 kg) of phosphorus and potassium were applied as basal during land preparation. The remaining dose (20 kg) of nitrogen was top-dressed one month after sowing. The growth of crop during the early stage was slow hence, weeding was necessary up to two months after sowing or up to a crop height of 40 cm. Coix is a tillering crop and all its tillers bear inflorescences two to three months after sowing (Fig. 8).
Pests And Diseases
Coi x also has a few diseases and pests. Among the diseases, the leaf blight caused by Bipolaris coicis is the most devastating disease of coix (Jansen 2006). In addition, the diseases like rust caused by Puccinia and smut by Ustilago and Tilletia have also been reported (Corke and Huang 2016). The Job’s tears smut is caused by Ustilago coicis which infects the ovary of flowers and leaves and is known to cause damage to crops in India, Thailand and China (Titatarn et al. 1983; Zhang et al. 2013). The rust disease is caused by Puccinia operta and leaf spot by Phyllachora coicis to the coix in Iran (Ahmadpour et al. 2013). Recently, the stem rot diseases of Coix lacrymajobi caused by Fusarium incarnatum were reported in China (Hou et al. 2022). Besides, the southern rice black-streaked dwarf virus (Pu et al. 2012) and maize chlorotic dwarf virus are viral diseases and the coix also hosts some bacterial diseases like leaf scald of sugarcane caused by Xanthomonas albilineans and gumming disease of sugarcane by Xanthonomas axonopodis pv. vasculorum (Hayward, 1993). Further, Coix is also attacked by number of insect pests such as Asian corn borer Ostrinia furnacalis, Asiatic rice borer Chilo suppressalis and rice skipper Pelopidas mathias (Ahmadpour et al. 2013; Kalaisekar et al. 2017). The infestation of root-knot nematode, Meloidogyne incognita has also been reported in Coix plants (Duke 1983). Recently, the invasive fall armyworm Spodoptera frugiperda was found ingesting Coix in China (Zhou and Yang 2019).
In our study, no disease incidence was observed on Coix plants. However, we recorded the infestation of fall armyworm, Spodoptera frugiperda (J.E. Smith) on Coix at the farms of ICAR-Research Complex for NEH, Mizoram Centre. The damage incidence of fall armyworm ranged between 7.23 to 10.86% in the vegetative growth stages (45 to 75 days crop) from the last week of August to the end of September. Fall armyworm larvae were found feeding on the leaves and leaf whorl of the Coix plants (Fig. 9). To the best of our knowledge, this is the first report of fall armyworm occurrence on Coix from India. However, fall armyworm infest more than 350 host plants including economically important cultivated grasses such as maize, rice, sorghum, sugarcane and wheat but also other vegetable crops and cotton (Montezano et al. 2018). The occurrence of fall armyworm on Coix was reported in China after its invasion of Asia (Zhou and Yang 2019). Besides, the fall armyworm prefers to feed on maize, however, in the absence of a preferred host, it can feed on other host plants such as Coix due to its polyphagous nature.
Harvesting
Crop usually matures in 4–5 months from the date of sowing depending upon cultivars and climatic conditions of the region. For good harvests, there must be enough rain during the early growth stages and dry weather is required during grain setting (Corke et al. 2016). At maturity, whole plants are cut at the basal region like rice using a sickle or dao and kept for sun drying. The grain is separated from the dried plant by threshing. Large size local wooden mortar (Mizo: sum) and pestle (Mizo: suk) are used to crush the dried seeds to remove the shell. The yield of husked grain generally ranges from 2–4 t/ha having a hulling percentage of 30–50% and fodder yield from 25–30 t/ha. If the stubble is left in the field and new growth and fresh leaves can be excellent fodder. Several cuts are possible if cultivated for fodder purposes. Under humid conditions, it is better to store unhusked grain than husked grain. The seeds of wild types are highly demanded by Art and Culture Department and other cultural clubs to make traditional head gear and other traditional ornaments and are purchased at Rs. 400–500/kg seeds (4.90 to 6.16 USD).
Problems And Prospects Of Coix In Mizoram
Coi x cultivation is very sporadic in Mizoram and its importance has been decreasing day by day and replacing with high-yielding cereals like maize and rice etc. Due to poor market demand, difficulty in processing due to the presence of hard seed coat, lack of knowledge on processing, non-availability of suitable varieties, ignorance on crop value and lack of scientific cultivation practices resulted in poor attention of Mizo farmers towards its cultivation. There is a lack of systemic knowledge on processing and processed products of this crop. Although, Mizoram is blessed with rich biodiversity, collection and proper documentation of genetic diversity are yet to be explored. Coix is closely associated with the socio-cultural importance of the Mizo community. This crop is hardy towards pests and disease attacks and can be grown in a less fertile area where other crops are unable to perform well and require lesser care. It is highly nutritious and promising medicinal properties fetch more research attention. At the same time, there is an increasing demand for Coix grains and products among health-conscious developed countries as a ‘functional food’. Therefore, the collection of germplasm, conservation, evaluation and selection of best performing genotypes are important for improving yield and quality with higher harvest index, dual-purpose types, early maturity, synchronous maturity and lodging-resistant properties. Utilizing genetic resources will also guarantee great productivity and good adaptability to the hilly terrain of the Mizoram ecosystem.