Breed. Biotech. 2014 (March) 2(1):48~63 Online ISSN: 2287-9366 http://dx.doi.org/10.9787/PBB.2014.2.1.048 Print ISSN: 2287-9358 RESEARCH ARTICLE

Preliminary Characterization and Evaluation of Landraces of Indian Spinach ( spp. L.) for Agro-economic and Quality Traits

Medagam Thirupathi Reddy1*, Hameedunnisa Begum1, Neelam Sunil2, Pandravada Someswara Rao2, Natarajan Sivaraj2, Sashi Kumar3 1Vegetable Research Station, Dr. Y.S.R. Horticultural University, Rajendranagar, Hyderabad-500 030, Andhra Pradesh, 2National Bureau of Plant Genetic Resources Regional Station, Rajendranagar, Hyderabad-500 030, Andhra Pradesh, India 3College of Horticulture, Dr. Y. S. R. Horticultural University, Rajendranagar, Hyderabad-500 030, Andhra Pradesh, India

ABSTRACT Indian spinach (Basella spp. L.) is an underutilized and underexploited indigenous leafy vegetable which has high nutritional and medicinal value and extensively used in the sub-continent. Landrace germplasm is endowed with rich genetic variability for various yield and quality traits. A total of six accessions collected through an exploration during 2010 were pre-bred by selfing during the October-January cropping season in 2011. These landraces were evaluated in a randomized block design with four replications in June-September, 2012 at Vegetable Research Station, Dr. Y. S. R. Horticultural University, Rajendranagar to assess the genetic diversity, variability, heritability and genetic advance for yield and its components in the material. Multivariate analysis following Ward’s minimum variance-method revealed distinct clustering pattern. Analysis of variance revealed highly significant differences among all genotypes for all the studied traits indicating considerable variability among ecotypes for most of the measured parameters. There was significant variability for genetic potential of all genotypes for different traits under study. The highest variability at genotypic level was observed for stalk yield (73.95%) followed by leaf-stalk ratio (46.70%) and weight of tender shoot (41.25%). Low to high estimates of broad sense heritability were found in different traits. High estimates of heritability (>60%) coupled with high genetic advance as percent of mean (>20%) for petiole length, internodal length, weight of tender shoot, stalk yield, leaf-stalk ratio and harvest index revealed that most likely the heritability is due to additive gene effects and selection may be effective. Keywords Genetic advance, Genetic diversity, Genetic variability, Heritability, Indian spinach, Landraces, Quality, Yield

INTRODUCTION parts of India, tropical Asia and . It is gaining popularity in some of the tropical and temperate climates of Indian spinach ( L., 2n=48; Basella rubra Asia, America, Australia and Europe. Although the plant is L., 2n=44) is one of the rapidly growing tropical leafy a perennial climber (Roshan et al. 2012), it is cultivated as vegetables, also noted by regional names in different an annual leafy vegetable. It is an important and reliable regions in Asia viz., Ceylon spinach, Malabar spinach, saan source of income for marginal and small farmers and tribal choy (Chinese), mong toi (Vietnamese), alugbati (), folks in southern states of India. It is widely adapted to a pui (Bengali), remayong (Malay) belongs to the variety of soils and climates. It thrives well in tropical and family and order . It is native to subtropical climates (Grubben 1997). Being a warm season tropical Asia, probably originating from India or Indonesia. crop, it is extremely heat tolerant (Grubben and Denton The distribution of Basella extends from the tropical and 2004) and frost tender. It can thrive under conditions of subtropical regions mostly America, Africa, Madagascar moderate soil fertility and can grow in highly acid soils, but and south India to New Guinea. It is grown in almost all is quite responsive to nitrogen fertilizer. It is a minor leafy

Received March 11, 2014; Revised March 27, 2014; Accepted March 28, 2014; Published March 31, 2014 *Corresponding author Medagam Thirupathi Reddy, [email protected], Tel: 040-24018016, Fax: 040-24018016

Copyright ⓒ 2014 by the Korean Society of Breeding Science This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 49

vegetable and since it is generally grouped together with over many centuries adapted to various eco-geographical other greens. The plant is found to be versatile in properties zones in India and thus the landraces accumulated significant (Saroj et al. 2012). It can be used as leafy vegetable, amount of variability. Several landraces of Basella are ornamental, dye and medicine. Tender shoots with under commercial cultivation in south India catering to the succulent stem along with thick, semi-succulent and requirements of local markets. Heterogeneity in production mucilaginous leaves are used as leafy vegetable. It is and quality of crop is mainly due to lack of adequate efforts appreciated for its organoleptic characteristics (Lucas towards characterization and evaluation of sizeable germplasm 1988) and makes a tasty addition to salads, dips and meals. and identification of elite accessions. Germplasm is the The leaves are also used for making soups (Lucas 1988). It reservoir of genetic diversity. Germplasm collections has high economic, nutritional and medicinal values. This play a key role for genetic improvement of any crop. crop is a valuable income source for marginal and small Characterization and evaluation data of germplasm will farmers near the principal urban centers. It plays a vital role increase utilization of germplasm so that people can make in food and nutritional security particularly during the dry better use of biodiversity (Bioversity International 2007). periods (Maundu et al. 1999). It is rich in food value, Genetic improvement of crops for quantitative traits supplying minerals, , proteins, and requires reliable estimates of genetic diversity, genetic dietary fibre (Adeboye 1996). It is found to be a good variability, heritability and genetic advancement in respect source of calcium, iron, A, vitamin B9 and Vitamin to the breeding material that is presently at hand in order to C (Lucas 1988; Okigbo 1990; Palada and Chang 2003; plan an efficient breeding program (Dudley and Moll 1969; Grubben and Denton 2004; Prance and Nesbitt 2005). Ulloa et al. 2007; Chand et al. 2008). An accurate Basella alba has been used from a long time back for the estimation of genetic diversity can be invaluable in the treatment of many diseases like dysentery, diarrhea, selection of diverse parental combinations to generate anemia and cancer (Roshan et al. 2012). The Ayurvedic segregating progenies with maximum genetic variability treatment in India used B. alba leaves and stems for and introgressing desirable traits from diverse or wild anticancer such as melanoma, leukemia and oral cancer germplasm into the available cultivars to broaden the (Premalatha and Rajgopal 2005). In India, it has been used genetic base (Ulloa et al. 2007). Genetic variability for for antipruritis and burn (Saikia et al. 2006), and has been agronomic traits is the key component of breeding used in Bangladesh for acne and freckle treatment (Akhter programmes for broadening the gene pool of crops. In view et al. 2008). Stems and leaves are used as mild laxative, of the above, collection, characterization and evaluation of diuretic and antipyretic (Chou 1997). Roots and leaves germplasm facilitate estimation and better utilization of have been used for the removal of after birth and stomach available diversity (Dudley and Moll 1969; Bioversity pains and for increasing milk production (Pascaline et al. International 2007; Ulloa et al. 2007; Chand et al. 2008). 2010). A decoction of the leaves is a good laxative for Until recently, research was concentrated mainly on pregnant women and children (Kirtikar and Basu 1975). major leafy vegetable species and very little attention has The boiled leaves along with sorghum flour are an effective been given to minor but indigenous leafy vegetable species. antiulcer agent (Dixit and Goyal 2011). The roots are used Basella is an indigenous minor leafy vegetable crop of in the treatment of diarrhea, the cooked leaves and stems increasing value but underutilized and underexploited. are used as laxatives (Larkcom 1991; Phillips and Rix Despite its multiple virtues, Basella crop is not included in 1995). The juice of leaves has been prescribed against the priority leafy vegetables for research in India. Until constipation especially for children and pregnant women recently no or little attention was paid to its genetic (Duke and Ayensu 1985). The flowers are used as an improvement. From the survey of literature, it is evident antidote to poisons and also as diuretic and febrifuge (Duke that genetic studies in Basella is limited, this trend is and Ayensu 1985). associated with little preference for this crop among Basella in the process of domestication and cultivation researchers in the world, often termed ‘orphan crop’. 50 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

Genetic studies in Basella revealed that phenotypic Genetic Resources Regional Station, Hyderabad in coefficient of variation was higher than genotypic coefficient collaboration with Vegetable Research Station, Dr. Y.S.R. of variation for all the traits studied, indicating environmental Horticultural University, Rajendranagar during 2010. influence on expression of these characters (Varalakshmi Following random sampling strategy, six landraces of and Devaraju 2010). Moderate heritability along with high Basella were collected from two districts of Andhra genetic advance was recorded for leaf weight and total Pradesh and one of Orissa from different locations (Table plant weight, indicating the presence of additive gene 1). DIVA-GIS version 7.5.0, free downloadable software effects. Higher plant weight was found to be significantly (Hijmans et al. 2012) was used to map the collection sites and positively associated with branch number, leaf number, of Basella germplasm from Andhra Pradesh and Orissa leaf weight and stem weight. Thus, for yield improvement (Fig. 1). in Basella, emphasis may be laid on indirect selection through leaf characters like leaf number, leaf length and Pre-breeding and initial seed increase leaf weight. Hence, selection can be employed for Basella being a short day plant which requires less than improvement of these characters in Basella (Varalakshmi 13 hours to induce flowering, was grown during rabi and Devaraju 2010). (October-January) season in 2011 at the Experimental The objectives of the study were to estimate the amount Farm, Vegetable Research Station, Rajendranagar for of genetic variability available in the Indian spinach maintenance and initial seed increase. Each of the six accessions and to estimate heritability and genetic advance landraces collected were raised in triple-row plots of among agronomic traits for selection. 3.0×1.8 m with inter-row spacing of 0.60 m and intra-row spacing of 0.30 m maintaining a population of 30 per genotype. Basella although autogamous, the flowering MATERIALS AND METHODS shoots were bagged with muslin cloth to prevent natural outcrossing. The fully ripe selfed fruits were harvested and Exploration survey and germplasm collection dried. A survey was conducted by the National Bureau of Plant

Table 1. Passport data of the various landraces of Indian spinach. Collection site Accession Collector Common Botanical Vernacular Geo-reference ID No. name name name Latitude Longitude Altitude Village Mandal District State (°N) (°E) (m) Indian Basella Mattu Andhra RNIS-1 SNTV-25 18.77 84.41 35.05 Peddaveedhi Palasa Srikakulam Spinach alba L. bacchali Pradesh Indian Basella Mattu Andhra RNIS-2 SNTV-38 18.36 83.87 28.65 Bavajipeta Srikakulam Srikakulam Spinach alba L. bacchali Pradesh Indian Basella Mattu Andhra RNIS-3 SNTV-51 18.31 83.57 66.14 Chipurupalli Chipurupalli Vizianagaram Spinach alba L. bacchalli Pradesh Indian Basella Erra Patha Andhra RNIS-4 SNTV-68 18.59 83.37 131.37 Bobbili Vizianagaram Spinach rubra L. bacchali Bobbili Pradesh Indian Basella RNIS-5 SNTV-79 Bhaji 18.58 82.91 1009.80 Malliput Pottangi Koraput Orissa Spinach alba L. Indian Basella RNIS-6 SNTV-86 Bhaji 18.63 82.59 605.33 Lamtaput Lamtaput Koraput Orissa Spinach alba L. RNIS= Rajendranagar Indian spinach Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 51

Experimental material, design and crop management carried out to safeguard the crop from pests and diseases. A total of six landraces of Basella (RNIS-1 to RNIS-6) thus collected and pre-bred were utilized for the present Recording of biometric and qualitative data study (Table 1). These six landraces were evaluated in a The plants of Basella are subjected to periodical randomized block design with four replications at the harvesting following shoot pickings at regular intervals as Experimental Farm (latitude 17.19oN and longitude practiced by farmers. All the landraces were harvested 79.23oE, and altitude 542.6 m) of the Vegetable Research simultaneously with first shoot picking at 45 days after Station, Rajendranagar under irrigated conditions during sowing and subsequent harvests at 30 days interval over a kharif (June-September) 2012. Since long days of ≥13 period of 3 months. Observations on the agro-economic hours precludes flowering in Basella, landraces were characters like leaf length (cm), leaf width (cm), leaf evaluated for leafy vegetable production during long days weight (g), petiole length (cm), length of tender shoot (cm) of kharif (June-September) for yield and quality traits. and internodal length (cm) were recorded from the random Each block consisted of 18 rows with 3 rows per genotype. sample of 10 tender shoots selected from ten randomly Each genotype was raised in a triple row plot of 3.0 m selected competitive plants, while the observations on length and 1.8 m width. In each replication, a plant other characters like leaf yield (g/shoot), stalk yield population of 10 plants per row and 30 plants per plot and (g/shoot), total biomass (g/plant), leaf-stalk ratio and genotype was maintained. Two seeds per hill were dibbled harvest index were recorded on whole plot basis in each with an intra-row spacing of 0.30 m and an inter-row entry in each replication. Young tender shoots consisting of spacing of 0.60 m. Thereafter, hills were thinned to one 10 intact leaves were harvested by clipping the stem with plant at two weeks after sowing. Recommended package of sharp knife. At first harvest in each treatment and practices was followed to raise a successful crop. All replication, ten tender shoots with 10 leaves from the agronomic practices were maintained for a whole duration growing tip were utilized to record the biometric data on of the experiment. Regular plant protection measures were leaf lamina length (cm), leaf lamina width (cm), leaf weight

Fig. 1. DIVA-GIS mapping of collection sites of landraces of Indian spinach from Andhra Pradesh and Orissa. 52 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

(g), petiole length (cm), length of tender shoots (cm), and high (>20.00%) as suggested by Sivasubramanian and internodal length (cm), weight of tender shoots, leaf Menon (1973). Heritability in the broad sense was derived yield/shoot (g) and stalk yield/shoot (g). These sampled based on the formula given by Allard (1960) and Burton tender shoots were stripped off leaves to facilitate and de Vane (1953). The heritability values were classified recording of data on separated stalks and leaves. Leaf as low (<30.00%), moderate (30.00-60.00%) and high length (cm) was determined by measuring all 10 leaves (>60.00%) as suggested by Johnson et al. (1955). The stripped off from the individual shoots from the leaf base to estimates of genetic advance (GA) at 5% selection intensity the tip with a measuring scale. Leaf width (cm) was (2.06) and genetic advance as percent of mean were determined by measuring all 10 leaves stripped off from the obtained using procedure given by Allard (1960). The individual shoots at the widest point of the leaf blade with estimates of genetic advance and genetic advance as a measuring scale. Average leaf weight (g) was determined percent of mean were classified as low (<10.00%), moderate by weighing all 10 leaves stripped off from the individual (10.00-20.00%) and high (>20.00%) as suggested by shoots in a digital analytical balance (±0.001 g). Length of Johnson et al. (1955). Hierarchical clustering (Fig. 3) was tender shoot consisting of 10 leaves from the growing tip carried out following Ward’s minimum variance method was measured in centimeters with a measuring scale. (Ward 1963). Length of tender shoot (cm) was divided with the number of nodes on the tender shoot to get internodal length (cm). Weight of 10 leaves stripped off from the tender shoots was RESULTS measured in a digital analytical balance (±0.001 g) to arrive at leaf yield per shoot (g). Weight of 10 tender shoots Characterization of landraces excluding leaves was measured in a digital analytical Germplasm characterization was centered primarily on balance (±0.001 g) to arrive at stalk yield per shoot (g). two major species, Basella alba L. and Basella rubra L. Of Data from each harvest were combined at the end of the the six landraces under study, RNIS-1 RNIS-2 RNIS-3 growing season to estimate the total shoot yield per plot and RNIS-5 and RNIS-6 with green leaf margin colour were total shoot yield per plant. Harvest index was taken as the recognized as Basella alba L., while RNIS-4 with red leaf ratio of leaf weight to total above-ground weight. The margin colour was recognized as Basella rubra L. Germplasm landraces were also characterized for 10 qualitative traits characterization was centered secondarily on qualitative like growth habit, stem colour, stem shape, leaf colour, leaf traits. Six landraces of Indian spinach characterized in this margin, leaf margin colour, leaf apex, petiole colour and study showed a broad variation for most of the qualitative flower colour on single plant basis. traits under study (Table 2). The variation in growth habit, stem colour, stem shape, leaf colour, leaf shape, leaf Statistical analysis of data margin colour, petiole colour and flower colour, were The data thus recorded were subjected to analysis of easily recognizable with visual appraisal in the material variance (Steel and Torrie 1980). The mean performance of (Fig. 2). The comparative view of plants, tender shoots and each individual genotype was employed in the statistical leaves of six landraces is depicted in Fig. 2. Plants of six analysis and to estimate major genetic components using landraces displayed great diversity in their growth habit SAS Enterprise Guide Version 4.2 (SAS Institute Inc (twining/ twining but initially bushy/ procumbent). Of the 2009). The genotypic coefficient of variation (GCV) and six landraces, the landraces RNIS-1 and RNIS-4 had phenotypic coefficient of variation (PCV) were computed twining growth habit. The landraces RNIS-2 and RNIS-3 by adopting the method of Burton and de Vane (1953) and had twining but initially bushy growth habit. The landraces Burton (1952). The phenotypic coefficient of variation RNIS-5 and RNIS-6 had procumbent growth habit. Stem (PCV) and genotypic coefficient of variation (GCV) values was red coloured in RNIS-1, RNIS-4 and RNIS-5 and were classified as low (<10.00%), moderate (10.00-20.00%) green coloured in RNIS-2, RNIS-3 and RNIS-6. Stems Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 53

were round (RNIS-1, RNIS-2 and RNIS-3) to angular margin (entire) and leaf apex (acuminate) among landraces. (RNIS-4, RNIS-5 and RNIS-6) in shape. Leaves were light Leaf margin was green in almost all accessions except green (RNIS-2, RNIS-3 and RNIS-6) to dark green RNIS-4 (red). Petioles are green in almost all accessions (RNIS-1, RNIS-4 and RNIS-5) in colour. Leaves were oval except RNIS-4 (red). Flowers are light pink (RNIS-2 and (RNIS-4 and RNIS-5) to ovate ((RNIS-1, RNIS-2, RNIS-3 RNIS-6) to pink (RNIS-1, RNIS-3, RNIS-4 and RNIS-5) in and RNIS-6) in shape. There was no variation in leaf colour. There exists no relationship between the stem

Fig. 2. Phenotypic variability in the plants (top row), tender shoots (middle row) and leaves (bottom row) of landraces of Indian spinach.

Table 2. Qualitative traits of six landraces of Indian spinach.

Landrace Character RNIS-1 RNIS-2 RNIS-3 RNIS-4 RNIS-5 RNIS-6 Twining Twining Growth habit Twining Twining Procumbent Procumbent (Initially bushy) (Initially bushy) Stem colour Red Green Green Dark red Light red Green Stem shape Round Round Round Angular (3) Angular (3) Angular (3) Leaf colour Dark green Light green Light green Dark green Dark green Light green Leaf shape Ovate Ovate Ovate Oval Oval Ovate Leaf margin Entire Entire Entire Entire Entire Entire Leaf margin colour Green Green Green Red Green Green Leaf apex Acuminate Acuminate Acuminate Acuminate Acuminate Acuminate Petiole colour Green Green Green Red Green Green Flower colour Pink Light pink Pink Pink Pink Light pink RNIS= Rajendranagar Indian spinach 54 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

colour of the accessions and the leaf colour as well as the leaf length, length of tender shoot, leaf yield and total petiole colour (Fig. 2). biomass. No significant differences were observed within replications for almost all traits except harvest index. Evaluation of landraces Analysis of variance Mean performance From the analysis of variance (Table 3), it is evident that The ranges of mean values revealed sufficient variation highly significant differences among six landraces were for all the traits under study (Table 4). Wide range of observed for majority of the characters under study except variation was observed in most of the quantitative characters.

Table 3. Analysis of variance for various agro-economic traits in Indian spinach.

Mean squares Character Replications Genotypes Error (3) (5) (10) Leaf length (cm) 0.736 2.014 0.658 Leaf width (cm) 0.101 1.183* 0.296 Leaf weight (g) 0.005 0.014** 0.002 Petiole length (cm) 0.003 0.528** 0.005 Length of tender shoot (cm) 16.243 24.423 12.388 Internodal length (cm) 0.305 2.780** 0.099 Weight of tender shoot (g) 2.768 114.353** 5.281 Leaf yield (g/shoot) 2.802 2.747 0.984 Stalk yield (g/shoot) 3.239 69.161** 5.205 Total biomass (g/plant) 25126.050 45971.820 22929.390 Leaf-stalk ratio 0.368 2.544** 0.187 Harvest index 0.060* 0.069** 0.009 *, **Significant at 5 and1 percent level, respectively Values in parentheses denote degrees of freedom

Table 4. Mean performance of six landraces for various agro-economic traits in Indian spinach.

Length Weight Leaf Leaf Leaf Petiole of Internodal of Total Leaf yield Stalk yield Leaf-stalk Harvest Landrace length width weight length tender length tender biomass (g/shoot) (g/shoot) ratio index (cm) (cm) (g) (cm) shoot (cm) shoot (g/plant) (cm) (g) RNIS-1 5.69 3.62 0.76 1.07 38.55 2.21 11.54 7.81 4.31 547.67 1.83 0.68 RNIS-2 5.26 3.65 0.77 1.04 41.43 2.76 11.31 8.05 3.17 532.33 2.62 0.73 RNIS-3 4.16 2.91 0.78 1.03 43.31 2.84 10.48 7.10 2.78 569.67 2.59 0.69 RNIS-4 4.73 2.95 0.74 1.00 46.79 2.80 10.47 7.45 2.77 473.33 2.77 0.73 RNIS-5 6.37 3.89 0.82 1.82 45.00 3.95 18.21 8.55 10.39 495.67 0.83 0.47 RNIS-6 5.99 4.54 0.93 1.87 42.78 4.82 25.71 9.79 14.04 814.67 0.75 0.38 S. Em 0.43 0.29 0.03 0.04 1.86 0.17 1.21 0.52 1.20 79.80 0.23 0.05 CD (5%) NS 0.99 0.09 0.13 NS 0.57 4.18 NS 4.15 NS 0.79 0.17 RNIS= Rajendranagar Indian spinach Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 55

In the materials used in this study, maximum range of genotypes from Andhra Pradesh state were grouped in one variability (Table 4) was observed for total biomass cluster, but one out of two genotypes from Orissa state was (473.33 to 814.67 g/plant) followed by weight of tender grouped in another cluster. shoot (10.47 to 25.71 g) and stalk yield (2.77 to 14.04 g/shoot). Of the six landraces, there was considerable Genetic variability analysis variation in shoot and leaf characters, which determine In general, phenotypic variances were higher than the external quality of tender shoots of Basella. Leaf size in corresponding genotypic variances for all the characters terms of lamina length was minimum in RNIS-3 (4.16 cm) under study (Table 5). The phenotypic variance was and maximum in RNIS-5 (6.37 cm) and in terms of lamina highest for total biomass (30610.20) followed by weight of width was minimum in RNIS-3 (2.91 cm) and maximum in tender shoot (41.64) and stalk yield (26.52). Similarly, the RNIS-6 (4.54 cm). RNIS-2 and RNIS-4 had highest harvest genotypic variance was also highest for total biomass index (0.73). RNIS-4 had highest leaf-stalk ratio (2.77) (7680.81) followed by weight of tender shoot (36.36) and followed by RNIS-2 (2.62). stalk yield (21.32). The degree of variability shown by different parameters can be judged by the magnitude of Genetic diversity analysis GCV and PCV. GCV showed that the extent of genetic Following Ward’s minimum variance method, 6 variability in the population ranged from 4.66 (length of landraces of Basella were grouped into two clusters as tender shoot) to 73.95 (stalk yield). PCV showed that the evident from the dendrogram (Fig. 3). Cluster I was the extent of genetic variability in the population ranged from largest with 5 genotypes, while cluster II was the smallest 9.42 (length of tender shoot) to 82.49 (stalk yield). The with only one genotype. Cluster I was subdivided into two estimates of PCV (Table 5) were highest for stalk yield sub clusters namely sub cluster IA with three genotypes (82.49%) followed by leaf-stalk ratio (51.97%) and weight and sub cluster IB with two genotypes. All of the four of tender shoot (44.14%), while lowest for length of tender

Fig. 3. Ward’s minimum variance dendrogram showing clustering of landraces of Indian spinach. 56 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

shoot (9.42%) followed by leaf weight (9.80%) and leaf The estimates of heritability (Table 5) were of high yield (15.43%). The estimates of GCV (Table 5) were magnitude (>60%) for leaf weight, petiole length, highest for stalk yield (73.95%) followed by leaf-stalk ratio internodal length, weight of tender shoot, stalk yield, (46.70%) and weight of tender shoot (41.25%), while leaf-stalk ratio and harvest index, of moderate magnitude lowest estimates were found for length of tender shoot (30-60%) for leaf length, leaf width and leaf yield and of (4.66%) followed by leaf weight (7.74%) and leaf yield low magnitude (<30%) for length of tender shoot and total (9.44%). The estimates of PCV (Table 5) were of high biomass. The estimates of genetic advance as percent of magnitude (>20%) for leaf width, petiole length, internodal mean (Table 5) were of high magnitude (>20%) for petiole length, weight of tender shoot, stalk yield, total biomass, length, internodal length, weight of tender shoot, stalk leaf-stalk ratio and harvest index, of moderate magnitude yield, leaf-stalk ratio and harvest index, of moderate (10-20%) for leaf length and leaf yield and of low magnitude (10-20%) for leaf length, leaf weight, leaf yield magnitude (<10.00%) for leaf weight and length of tender and total biomass and of low magnitude (<10%) for length shoot. The estimates of GCV (Table 5) were of high of tender shoot. High estimates of heritability (>60%) magnitude (>20%) for petiole length, internodal length, coupled with high genetic advance as percent of mean weight of tender shoot, stalk yield, leaf-stalk ratio and (>20%) were observed for petiole length, internodal length, harvest index, of moderate magnitude (10-20%) for leaf weight of tender shoot, stalk yield, leaf-stalk ratio and length, leaf width and total biomass and of low magnitude harvest index. The characters like petiole length, internodal (<10.00%) for leaf weight, length of tender shoot and leaf length, weight of tender shoot, stalk yield, leaf-stalk ratio yield. In general, the magnitude of phenotypic coefficients and harvest index recorded high genetic advance as percent of variation (PCV) was higher than the corresponding of mean coupled with high heritability estimates. genotypic coefficients of variation (GCV) for all the twelve Overall, the analysis of variance revealed considerable characters under study (Table 5). The magnitudinal variability among the landraces for the majority of the leaf differences between the estimates of GCV and PCV were yield and its components under study. Further, the results highest for total biomass (15.26) followed by leaf length of the genetic parameters also indicated considerable (7.11) and leaf width (6.26). variability in the material. There was considerable variation

Table 5. Genetic parameters for various agro-economic traits in Indian spinach.

Coefficient of variation Genetic Variance Genetic (%) Heritability advance as Character advance (%) percent of (5%) Phenotypic Genotypic Phenotypic Genotypic mean Leaf length (cm) 1.11 0.45 19.64 12.53 41.00 0.88 16.47 Leaf width (cm) 0.59 0.30 21.36 15.10 50.00 0.79 21.98 Leaf weight (g) 0.01 0.00 9.80 7.74 62.00 0.10 12.59 Petiole length (cm) 0.18 0.17 32.46 31.96 97.00 0.85 64.84 Length of tender shoot (cm) 16.40 4.01 9.42 4.66 25.00 2.04 4.75 Internodal length (cm) 0.99 0.89 30.85 29.27 90.00 1.85 57.22 Weight of tender shoot (g) 41.64 36.36 44.14 41.25 87.00 11.61 79.40 Leaf yield (g/shoot) 1.57 0.59 15.43 9.44 37.00 0.97 11.89 Stalk yield (g/shoot) 26.52 21.32 82.49 73.95 80.00 8.53 136.58 Total biomass (g/plant) 30610.20 7680.81 30.58 15.32 25.00 90.44 15.80 Leaf-stalk ratio 0.97 0.79 51.97 46.70 81.00 1.64 86.43 Harvest index 0.03 0.02 27.75 22.99 69.00 0.24 39.22 Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 57

in the coefficient of variation, heritability and genetic conservation (Rubenstein and Heisey 2003). This enables advance of various quantitative traits under study offering the identification of unique genotypes essential for curators ample scope for selection. There was considerable divergence of gene banks. Hence, characterization of newly collected among the landraces as evident from the genetic diversity germplasm is a necessary requirement for crop improvement, analysis. use and conservation of plant genetic resources. Basella alba L. features green-stems and deep-green leaves, while Basella rubra L. is known for its purplish- DISCUSSION stem and deep-green leaves with pink veins. The striking morphological difference between Basella alba L. and One of the main reasons for the under-utilization of Basella rubra L. is the colour of the edges of the leaves and germplasm, according to curators, breeders and other users the stem. Accessions with green leaf margin colour were of plant genetic resources, is the lack of adequate passport, recognized as Basella alba L., while RNIS-4 with red leaf characterization and evaluation data: people cannot use margin colour was recognized as Basella rubra L. The genetic resources that lack essential information. In addition, accessions belonging to these two species were identified such information is necessary for proper management of with the help of Economic Botanists from National Bureau the resources in the gene banks by gene bank managers. of Plant Genetic Resources Regional Station, Rajendranagar. Therefore, the accurate documentation of information Botanical characterization primarily revealed that there about the origin, characterization and performance of were two species Basella alba L. (RNIS-1 RNIS-2 RNIS-3 germplasm is essential for effective conservation and use. RNIS-5) and Basella rubra L. (RNIS-6) under cultivation Characterization and evaluation are important to provide in South India. The purple form, B. rubra L., is less breeders, growers and consumers, with information about preferred to the green type B. alba L., regardless of their certain properties of newly collected genotypes. nutritional value (Edema and Fakorede 1978). Six landraces of Indian spinach characterized in this Characterization of landraces study showed a broad variation for most of the qualitative Characterization of genetic resources refers to the traits (Table 2), which allows for the identification of process by which accessions are identified, differentiated promising landraces of Indian spinach. The apparent or distinguished according to their character or quality variation in qualitative traits could probably be due to the (traits) (Merriam-Webster 1991). Characterization of each fact that the genotypes used in this study were the sample involves a careful description of the special indigenous landraces and hence distinctly variable or characteristics that are inherited, easy to score and diverse with respect to qualitative traits. Landraces with expressed consistently in all environments (Rubenstein and twining but initially bushy growth habit are locally called Heisey 2003). Characterization on the basis of qualitative as ‘mattu bacchali’, while those with twining or procumbent traits provides information on diversity within and between growth habit are locally called as ‘teega bacchali’ in genotypes. Since most of the traits recorded during Andhra Pradesh, India. Twining growth habit is the characterization are those that can be seen, the person commonest growth habit. Twining plant type is advantageous responsible for managing the germplasm material is best to perennial leaf production since it would allow maximum placed to carry out the work of documenting these and uniform exposure or distribution of all leaves and other characteristics (De Vicente et al. 2005). Many of the vegetative parts for better interception of sunlight. This characteristics that are recorded on individual accessions would result in an increase in dry matter production and a can serve as diagnostic descriptors for the accessions subsequent increase in yield. Moreover, there is less chance (Rubenstein and Heisey 2003). Such diagnostic characters of tender shoots and leaves touching the ground or soil help gene bank curators keep track of an accession and thereby causing leaf rot as in case of procumbent growth check for the genetic integrity over a number of years of habit. The difference between the procumbent and 58 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

circumnutating stem forms was quite distinct particularly teams that include breeders, entomologists, pathologists in terms of biomass since the circumnutating form showed and agronomists (De Vicente 2005). A successful variety reduced laminar area and weight but with long internodes in any vegetable crop must meet minimal criteria for and minor stem circumference, which is otherwise numerous traits that are potentially valued in the markets. perfectly suited for circumnutation. Of the six landraces, Superiority for multiple ‘yield’ and ‘quality’ traits is the landraces RNIS-1 and RNIS-4 with twining growth essential for economic sustainability of a variety. Consumer habit could be exploited for commercial cultivation on purchases of Basella are enthused based on ‘quality’ traits supports. The landraces with twining growth habit are inherent to the leaves such as size, shape and color. Thus, often grown on supports like stakes or trellises or pandals. yield is partly about producing tonnage, but also about the On reaching a height of 30 cm, the plant is usually proportion of the crop that can be harvested and brought to supported by sticks (Lucas 1988). The three types of market in a condition and at a price acceptable to the staking-individual, pyramidal, and trellising could be consumer. adopted, but for optimum yield, a trellis or stick table of From the analysis of variance (Table 3), it is evident that about 30 cm above the surface of the bed should be highly significant differences among the landraces were constructed. However, for individual staking to be observed for majority of the characters under study effective, the stakes must be stout and at least 2 m tall. The indicating presence of sufficient amount of variability in landraces RNIS-2 and RNIS-3 with twining but initially the germplasm under study. Such wide variation indicated bushy growth habit could be exploited for pot culture for the scope for improving the population for these characters. urban vegetable cultivation. The landraces RNIS-5 and Non-significant differences within replications for almost RNIS-6 with procumbent growth habit could be exploited all traits except harvest index suggested that the landraces for ground culture. The plants of twining growth habit are have enough variability for almost all the traits to carry out exceedingly vigorous and do better with support than those further genotypic studies. Generally, the significant of procumbent growth habit when sprawling on the ground differences (P<0.01) revealed among the aforementioned to keep the foliage clean. morpho-agronomic traits may be due to environmental influences on the genotypes as well as differences in the Evaluation of landraces genetic potential of the different landraces of Indian Evaluation goes deeper than characterization. It may spinach. This indicates the role of environmental factors as require special biometrical techniques and usually include well as differences in the genetic makeup of different agronomic performance, yield and biotic and abiotic landraces in leaf yield determination of Indian spinach. The stresses, such as drought or pest. These traits are important non-significant differences observed in the landraces for to plant breeders and researchers in crop improvement. leaf length, length of tender shoot, leaf yield and total Such evaluation may also use DNA-based methods to biomass indicates that the genetic components of the analyze a plant's genetic diversity (Rubenstein and Heisey genetic material are intact. It follows, therefore, that any 2003; De Vicente et al. 2005). The evaluation descriptors, improvement sort must be directed at the other characters although contributing to some extent to identifying an except leaf length, length of tender shoot, leaf yield and accession, are more interesting than characterization total biomass. descriptors because of their value in crop improvement. In In conventional breeding programs, the choice of general, effective evaluation is possible when there is close parents, in general, is based on the general principle that the institutional and personal interaction between curators and parents under selection should have a high mean breeders or other crop improvement scientists, and where performance for the desirable traits. Hence, the breeders breeding objectives are reflected in evaluation programs are in absolute need of desirably high or low mean value (Rubenstein and Heisey 2003; De Vicente et al. 2005). depending upon the character, which is considered as a Evaluation is primarily carried out by users, in multidisciplinary main criterion for effective selection forever. In Basella, of Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 59

the twelve quantitative traits under study, high mean values practice in southern parts of India (Saroj et al. 2012). The are desirable for leaf length, width, weight and yield, succulent young leaves and stems are consumed as a weight of tender shoot, total biomass, leaf-stalk ratio and vegetable in this crop. Accordingly, Basella is harvested by harvest index, while low mean values are desirable for periodical shoot pickings. In general, bunches of fresh petiole length, internodal length and stalk yield. Due young tender shoots are marketed for fresh consumption of reduction in internodal length adds the more number of leaf Basella. For marketing, ten young tender shoots of 30 cm production points in plant. High leaf-stalk ratio and harvest length are bunched and tied with a rope or thread. In quality index are the key aspects. Harvest index is not directly a breeding, the issue that leafy vegetable breeder encounters yield contributing trait but is considered important parameter is how to assess quality. Ideally, the way it is assessed in the for genetic improvement of genotypes. After analyzing the breeding program should reflect the quality that the mean performance of 6 different Indian spinach landraces consumer will perceive. Because the consumers are the (Table 4), it is concluded that the two landraces RNIS-2 and ultimate users of the produce of any crop and possess RNIS-4 remained superior in terms of leaf-stalk ratio and valuable traditional knowledge, due consideration must be harvest index. It is, therefore, recommended that these given to involve consumers’ views and expectations at genotypes can be brought forward to evaluate the yield some point during any evaluation program. Accordingly, constituency in various ecological zones across the Andhra during exploration survey consumer groups were also Pradesh State. Upon assessing yield stability through interviewed at local vegetable markets to determine the multi-location trials, these genotypes may be used for external quality traits that they consider as acceptable. The large-scale cultivation if found suitable. Evaluation of qualitative characters based on which most consumers eleven germplasm lines of the Indian spinach (Basella) determine the quality of Basella were stem length (short/ revealed maximum leaf weight per plant in IIHR-1 (160.5 long), stem straightness (straight/curved), stem thickness g), followed by IIHR-18 (111.6 g) and IIHR-3 (98.3 g) (thin or thick) stem shape (oval or ovate) and branch color (Varalakshmi and Devaraju 2010). (green or purple), leaf size (small, medium or large), leaf shape (oval or ovate), leaf color (green or light green or Quality of landraces dark green or red). In local markets, shoot quality plays an A plant breeder is usually interested in accessions which important role in marketability. Shoot quality is mainly possess some desired quality traits for incorporation in related to the stem straightness, length, shape and colour breeding programs. Quality is often defined from either a and intact leaf size, shape and colour. The tender shoots product orientation or a consumer orientation. Quality is a intended for market must mostly be straight, short, thin and measure of how much the end-user values a product. The green. Leaf colour is commonly used measure of leaf term quality implies the degree of excellence of a product quality. A specific greenness and leaf size are also required, or its suitability for a particular use. Quality is a human but they are often arbitrarily indicated. Size is another construct comprising many properties or characteristics. factor that affects market acceptance of the leaf that is Leaf quality plays an important role in marketability of produced. External amount and intensity of surface colour leafy vegetables. What can leafy vegetable growers do to on a leaf of any cultivar is of prime importance to its appeal manage the quality of their produce? One of the first steps in the marketplace. For example, light green colored and leafy vegetable growers can take to ensure good quality is medium sized leaves are the most preferred quality traits. In variety selection. What can leafy vegetable breeders do to fact, the quality traits like greenness and size (length and assess the quality of their varieties? When new leafy width) of the leaves are associated with Basella genotype. vegetable germplasm have been collected, they should be In the surveyed local markets, local landraces with green evaluated in variety trials based on certain external leaf and round stalk possessing leaves with green/dark green characteristics to decide their quality. colour, oval/ovate shape, entire leaf margin, green leaf In Basella, both stem and leaves are used in culinary margin colour, acuminate leaf apex are acceptable/desirable 60 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63

qualitative traits of Indian spinach. A high level of influences, heritability of the characters and genetic advance variability was observed among the accessions in the that can be realized in practical breeding in evolving quality. On the whole, the results indicated a high level of varieties to various environmental conditions. Genetic variability among the accessions in the quality and the variability is essential in order to realize response to possibility to use these landraces to improve the agro- selection pressure. Perusal of data in Table 5 shows a economic and quality traits during breeding programs. considerable difference between PCV and GCV values for RNIS-2, RNIS-3 and RNIS-6 were among the landraces all the characters studied. This indicates presence of greater that had acceptable quality traits. environmental influence on expression of all these characters and selection may not be effective in improvement of Genetics of yield traits Basella. This also indicates that direct selection is not The determination of genetic diversity of accessions is effective in these characters and that heterosis breeding can extremely important in breeding of crop plants. Genetic be resorted to for further improvement. Similar observations divergence analysis estimates the extent of diversity were made by earlier researcher in Basella (Varalakshmi existed among selected genotypes. Precise information on and Devaraju 2010). Since genotypic coefficient of the nature and degree of genetic diversity helps the plant variation compares the relative amount of variability breeder in choosing the diverse parents for purposeful among attributes, it could, therefore, be deduced that stalk hybridization. The value of a germplasm collection yield, leaf-stalk ratio and weight of tender shoot had higher depends not only on the number of accessions it contains, amount of exploitable genetic variability among the but also upon the diversity present in those accessions. A attributes. It also signifies that there is greater potential for small set of accessions of Indian spinach was utilized for favorable advance in selection in these attributes when divergence analysis in the present study owing to the fact compared to others. High degree of genetic variability for that it is commercially cultivated in only few tracts of most of the characters in the present investigation offers a Andhra Pradesh and Orissa. Hence, it is indispensable to greater scope for effective selection. scrutinize and exploit this natural genetic diversity of In general, the magnitude of phenotypic coefficients of Basella for crop improvement. Ward (1963) describes a variation was higher than the corresponding genotypic class of hierarchical clustering methods including the coefficients of variation for all the twelve characters under minimum variance method. Ward's minimum variance study (Table 5) indicating environmental influence on method tends to join clusters with a small number of expression of these characters. However, the differences observations, and it is strongly biased toward producing between PCV and GCV were narrow indicating low clusters with roughly the same number of observations. It is environmental influence in the expression of these also very sensitive to outliers (Milligan 1980). Hierarchical characters. The magnitudinal differences between the clustering following Ward’s minimum variance method estimates of GCV and PCV were highest for total biomass revealed the presence of wide range of diversity in agro- (15.26) followed by leaf length (7.11) and leaf width economic traits offering a great scope for improvement of (6.26). Varalakshmi and Devaraju (2010) also observed Basella. These studies have shown that accessions from the relatively higher magnitude of PCV than GCV for leaf same geographical region may differ genetically as well as weight, stem weight, petiole length, leaf length, leaf phenotypically and also in adaptability. The pattern of breadth and total plant weight in Basella. Presence of this clustering showed more or less distinct relationship with high variability for the above parameters can form basis for the geographic origin of landraces. Geographical origin effective selection of superior lines in Basella. For other was found to be a good parameter of genetic divergence. characters, the PCV and GCV values were close to one Genetic variability analysis is helpful to know about the another, implying that genotype contributed more to the nature and extent of variability that can be attributed to expression of these characters than environment, different causes, sensitive nature of the crop to environmental suggesting greater possibilities of improvement through Preliminary Characterization and Evaluation of Landraces of Indian Spinach (Basella spp. L.) for Agro-economic and Quality Traits ∙ 61

selection. With the help of GCV alone, it is not possible to selection for the improvement of such characters may determine the extent of heritable variation. Thus the not be useful, because broad sense heritability is based estimates for heritability indicate effectiveness with which on total genetic variance which includes both fixable selection may be expected to exploit existing genetic (additive) and non-fixable (dominance and epistatic) variability. variances. Such high heritability values in yield and its The observed variability is a combined measure of related characters were also reported in Indian spinach genetic and environmental causes, whereas genetic (Varalakshmi and Devaraju 2010). The high heritability, variability alone is heritable. For improvement of leaf therefore, implies that these yield attributes are controlled yield in Basella direct selection is often misleading. The genetically, signifying high potential for improvement genetic variability is heritable from generation to through selection. The moderate estimates of heritability generation. Heritability and genetic advance is a useful for leaf length, leaf width and leaf yield indicating that tool for breeders in determining the direction and these characters are moderately influenced by environmental magnitude of selection. Yield and yield components, in effects and genetic improvement through selection will general, are polygenic in nature and are subjected to be moderately difficult due to masking effects of the different amount of non-heritable variation (Lush 1940). environment on the genotypic effects. Effectiveness of selection is dependent not only upon The estimates of heritability alone fail to indicate the the nature, extent and magnitude of genetic variability response to selection. Therefore, heritability estimates present in material but also on the extent to which it appear to be more meaningful when accompanied by is heritable. Heritability gives the information on the estimates of genetic advance and genetic advance as magnitude of inheritance of quantitative traits, while percentage over mean (Johnson et al. 1955). The estimates genetic advance will be helpful in formulating suitable of genetic advance as percent of mean (Table 5) were of selection procedures. Heritability in conjunction with high magnitude (>20%) for petiole length, internodal genetic advance has a greater role to play in determining length, weight of tender shoot, stalk yield, leaf-stalk ratio the effectiveness of selection of a character (Johnson et and harvest index, of moderate magnitude (10-20%) for al. 1955). leaf length, leaf weight, leaf yield and total biomass and of The ratio of genetic variance to the total variance i.e., low magnitude (<10%) for length of tender shoot. High phenotypic variance is known as heritability. Heritability estimates of heritability (>60%) coupled with high genetic in broad sense is the ratio of genotypic variance to total advance as percent of mean (>20%) for petiole length, variance in non-segregating population (Hanson et al. internodal length, weight of tender shoot, stalk yield, 1956). Heritability estimates gives a measure of leaf-stalk ratio and harvest index revealed that most likely transmission of characters from one generation to the the heritability is due to additive gene effects and selection next and the consistency in the performance of progeny may be effective. Such value of high heritability and high in succeeding generations and depends mainly on the genetic advance may be attributed to the action of additive magnitude of heritable portion of variation. Heritability genes (Panse 1957). The characters like petiole length, is the only component which is transmitted to the next internodal length, weight of tender shoot, stalk yield, generation. Heritability estimates of quantitative characters leaf-stalk ratio and harvest index recorded high genetic play an important role in expressing the reliability of advance as percent of mean coupled with high heritability variance value as a selection guideline to the plant estimates, indicating that these traits were under the strong breeder during the succeeding generations. High values influence of additive gene action, and hence simple selection of heritability for leaf weight, petiole length, internodal based on phenotypic performance of these traits would be length, weight of tender shoot, stalk yield, leaf-stalk ratio more effective. Similar kind of results in Basella was also and harvest index indicated that though the characters reported by earlier researcher (Varalakshmi and Devaraju are least influenced by the environmental effects, the 2010). Low heritability and low genetic advance as percent 62 ∙ Plant Breed. Biotech. 2014 (March) 2(1):48~63 of mean values were observed for the character length of REFERENCES tender shoot. This indicates that this character is highly influenced by environmental effects and selection would Adeboye OC. 1996. Proximate composition and nutrient be ineffective. Similar results were also reported by other analysis of six selected leaf vegetables of South West researcher in Basella (Varalakshmi and Devaraju 2010). Nigeria. Ife J. Agric. 18: 56. Akhter S, Abdul H, Shawkat IS, Swapan KS, Mohammad The morphological evaluation and characterization SHC, Sanjay SS. 2008. 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