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<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article" dtd-version="3.0" xml:lang="en">
   <front>
      <journal-meta>
         <journal-id journal-id-type="publisher-id">SJAR</journal-id>
         <journal-title-group>
            <journal-title>Spanish Journal of Agricultural Research</journal-title>
            <abbrev-journal-title>SJAR</abbrev-journal-title>
         </journal-title-group>
         <issn pub-type="epub">2171-9292</issn>
         <publisher>
            <publisher-name>Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria (INIA)</publisher-name>
         </publisher>
      </journal-meta>
      <article-meta>
         <article-id pub-id-type="publisher-id">10190</article-id>
         <article-id pub-id-type="doi">10.5424/sjar/2017152-10190</article-id>
         <article-categories>
            <subj-group subj-group-type="heading">
               <subject>Research article</subject>
            </subj-group>
         </article-categories>
         <title-group>
            <article-title>
               Assessment of combining ability for yield and quality components in hot pepper (
               <i>Capsicum annuum</i>
               L.)
            </article-title>
            <alt-title alt-title-type="running-head">Combining ability analysis in chilli for yield and quality</alt-title>
         </title-group>
         <contrib-group>
            <contrib contrib-type="author" corresp="yes">
               <name>
                  <surname>Rohini</surname>
                  <given-names>Narayanasamy</given-names>
                  <aff>Tamil Nadu Agricultural University, Horticultural College and Research Institute, Dept. Vegetable Crops, Periyakulam-625 604, Tamil Nadu, India.</aff>
               </name>
            </contrib>
            <contrib contrib-type="author" corresp="no">
               <name>
                  <surname>Lakshmanan</surname>
                  <given-names>Valliyappan</given-names>
                  <aff>Tamil Nadu Agricultural University, Horticultural College and Research Institute, Dept. Vegetable Crops, Periyakulam-625 604, Tamil Nadu, India.</aff>
               </name>
            </contrib>
            <contrib contrib-type="author" corresp="no">
               <name>
                  <surname>Saraladevi</surname>
                  <given-names>Desabandhu</given-names>
                  <aff>Tamil Nadu Agricultural University, Horticultural College and Research Institute, Dept. Vegetable Crops, Periyakulam-625 604, Tamil Nadu, India.</aff>
               </name>
            </contrib>
            <contrib contrib-type="author" corresp="no">
               <name>
                  <surname>Amalraj</surname>
                  <given-names>John J.</given-names>
                  <aff>Tamil Nadu Agricultural University, Dept. Plant Genetic Resource, Coimbatore-641003, Tamil Nadu, India</aff>
               </name>
            </contrib>
            <contrib contrib-type="author" corresp="no">
               <name>
                  <surname>Govindaraju</surname>
                  <given-names>Perumal</given-names>
                  <aff>Tamil Nadu Agricultural University, ADAC&amp;RI, Dept. Soil Science and Agricultural Chemistry, Trichy -620 009, Tamil Nadu, India</aff>
               </name>
            </contrib>
         </contrib-group>
         <author-notes>
            <corresp>
               should be addressed to Rohini Narayanasamy:
               <email xlink:href="rohizna@gmail.com">rohizna@gmail.com</email>
            </corresp>
         </author-notes>
         <pub-date pub-type="epub">
            <day>01</day>
            <month>07</month>
            <year>2017</year>
         </pub-date>
         <pub-date pub-type="collection">
            <year>2017</year>
         </pub-date>
         <volume>15</volume>
         <issue>2</issue>
         <elocation-id content-type="doi">10.5424/sjar/2017152-10190</elocation-id>
         <history>
            <date date-type="recibido">
               <day>01</day>
               <month>06</month>
               <year>2016</year>
            </date>
            <date date-type="aceptado">
               <day>05</day>
               <month>06</month>
               <year>2017</year>
            </date>
         </history>
         <permissions>
            <copyright-statement>© 2017 INIA</copyright-statement>
            <copyright-year>2017</copyright-year>
            <license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc/3.0/">
               <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution (CC-by) Spain 3.0 License.</license-p>
            </license>
         </permissions>
         <abstract id="abstract01">
            <title>Abstract</title>
            <p>TSix parental genotypes and their thirty F1 hybrids in a full diallel cross system with reciprocals, were used in the present study to estimate combining ability for five quantitative and five qualitative traits in hot pepper. The mean sum of squares due to general combining ability (GCA), specific combining ability (SCA) and reciprocal combining ability (RCA) were significant for all the characters which suggest that these parents could have performed differently in different combinations indicating interallelic interactions. All the traits studied exhibited greater SCA variance than GCA, suggesting that these traits are controlled by non-additive gene action. Analysis of the GCA effects of parents for various traits revealed that LCA625, K1 and PKM1 were the best general combiners for almost all the traits. Estimates of SCA effects showed that the hybrids LCA625 × K1, K1 × Arka Lohit and Pusa Jwala × PKM1 were found to be the best combiners for yield and its attributes. The hybrid K1 × Arka Lohit was the best reciprocal combiner for quality parameters based on its better mean performance and combining ability. These results suggest that commercial production of hot pepper hybrids and isolation of pure lines from these hybrids is a possible way to enhance the yield and fruit quality.</p>
         </abstract>
         <kwd-group>
            <title>Additional key words:</title>
            <kwd>chilli;</kwd>
            <kwd>growth;</kwd>
            <kwd>capsaicin;</kwd>
            <kwd>dry yield;</kwd>
            <kwd>GCA;</kwd>
            <kwd>SCA;</kwd>
            <kwd>RCA</kwd>
         </kwd-group>
         <funding-group>
            <funding-statement>The authors received no specific funding for this work</funding-statement>
         </funding-group>
      </article-meta>
      <notes>
         <p>
            <bold>Author´s contributions:</bold>
            Designed the study: LV, SD, JJA and GP. Data collection, field work and statistical analysis: RN. Technical and material support: LV and SD. Contributed analysis tools: JJA and GP. All authors read and approved the final manuscript.
         </p>
         <p>
            <bold>Competing interests:</bold>
            The authors have declared that no competing interests exist.
         </p>
         <p>
            <bold>Citation:</bold>
            Rohini, N.; Lakshmanan, V., Saraladevi, D., Amalraj, J. J.; Govindaraju, P. (2017). Assessment of combining ability for yield and quality components in hot pepper (
            <i>Capsicum annuum</i>
            L.). Spanish Journal of Agricultural Research, Volume 15, Issue 2, e0703.
            <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.5424/sjar/2017152-10190">https://doi.org/10.5424/sjar/2017152-10190</ext-link>
         </p>
      </notes>
   </front>
   <body>
      <sec id="S1">
         <title>Introduction</title>
         <p>
            Hot pepper (
            <italic>Capsicum annuum</italic>
            L.) is the universal vegetable cum spice of India. In India it is cultivated in all the states and union territories of the country. The most important states growing chilli are Andhra Pradesh, Orissa, Maharashtra, West Bengal, Rajasthan and Tamil Nadu. Andhra Pradesh alone commands 46% of the chilli production in India. As per the latest statistics, India produced 11,000,452 tonnes of dry chillies from an area of 9,036,028 hectares. No country in the world has so much area and production of chilli. Almost 90% of chilli production is consumed indigenously while only 10% is exported. Demand from the chilli powder manufacturing sector constitutes 30% of the total production in the country. Exports of chilies around 347,000 tonnes, which makes 29.20% of the total spices exported from India (
            <xref ref-type="bibr" rid="b23">Rohini, 2015</xref>
            ).
         </p>
         <p>
            Hot pepper has two important commercial qualities. Some varieties are famous for red colour because of the pigment capsanthin, others are known for biting pungency attributed by capsaicin. India is the only country rich in many varieties with different quality factors. In the world, both domestic consumption and export are maximum in India. As a product, oleoresin of hot pepper with low, medium or high pungency is also exported in large quantities (
            <xref ref-type="bibr" rid="b14">
               Manjula
               <italic>et al.</italic>
               , 2011
            </xref>
            ). Chilli powder is another important product of export. Indian chillies and its products are brought out by many countries,
            <italic>viz</italic>
            ., Sri Lanka, Bangladesh, South Korea and USA for dry chillies and USA, Germany, Japan, UK and France for oleoresin. India can supply chilli as in whole, crushed, powder or oleoresin forms in consistent colour and required pungency (
            <xref ref-type="bibr" rid="b28">
               Sharanakumar
               <italic>et al.,</italic>
               2011
            </xref>
            ).
         </p>
         <p>
            Hot peppers have been standing out by their increasing preference in the consumer market. Even though increase in cultivation and commercialization of hot pepper in the last few years (
            <xref ref-type="bibr" rid="b17">
               Nascimento
               <italic>et al.</italic>
               , 2014
            </xref>
            ) there is increasing demand in India for new cultivars that have higher yield, quality, pest and disease resistance, all of which can be achieved through breeding programmes.
         </p>
         <p>
            Diallel analysis is a biometrical tool that provides the estimates of genetic parameters regarding heterosis and combining ability. It gives additional information on presence or absence of epistasis, average degree of dominance and distribution of dominant and recessive genes in the parents (
            <xref ref-type="bibr" rid="b20">
               Rego
               <italic>et al.,</italic>
               2011
            </xref>
            ;
            <xref ref-type="bibr" rid="b17">
               Nascimento
               <italic>et al</italic>
               ., 2014
            </xref>
            ). Application of diallel techniques to self-pollinated crops for improving yield and quality may be appropriate (
            <xref ref-type="bibr" rid="b7">Griffing, 1956</xref>
            ). The concept of combining ability was developed by
            <xref ref-type="bibr" rid="b32">Sprague &amp; Tatum as early as 1942</xref>
            . According to them, the general combining ability (GCA) is the comparative ability of the lines or parents to combine with other lines. Specific combining ability (SCA) is the deviation in the performance of a specific cross from the performance expected on the basis of GCA effects of the parents involved in the cross. The expression of heterosis is highly associated with SCA of crosses.
         </p>
         <p>
            Study of combining ability is useful in predicting the performance of hybrid. Simultaneously the genetic distance also plays a major role in to assess the hybrid vigour component (
            <italic>e.g</italic>
            .
            <xref ref-type="bibr" rid="b21">
               Rodríguez
               <italic>et al</italic>
               ., 2003
            </xref>
            ,
            <xref ref-type="bibr" rid="b22">2008</xref>
            ). Hence, combining ability analysis is extensively used to study nature and magnitude of genotypic variability and facilitate correct choice of desirable parents in a hybrid development programme. The knowledge of gene effects on expression of any trait greatly helps the plant breeder in making to draw strategies of breeding population, selection method, type of variety and extent of testing. Combining ability analysis also gives a clear picture of elite parent which can be fitted as a general combiner over a series of cross combinations and in a specific cross to exploit heterosis.
         </p>
         <p>
            <xref ref-type="bibr" rid="b24">Roja &amp; Sprague (1952)</xref>
            and
            <xref ref-type="bibr" rid="b7">Griffing (1956)</xref>
            stated that the GCA includes the additive genetic portion, while SCA includes non-additive genetic portion of the total variation.
            <xref ref-type="bibr" rid="b8">Hayman (1957)</xref>
            found that in the absence of epistasis, GCA is composed of both additive and dominance while SCA involves mainly dominance effects. Extensive work has been done in chilli by various research workers on the combining ability of the exotic and indigenous genotypes for various characters.
            <xref ref-type="bibr" rid="b9">Jagadeesha &amp; Wali (2008)</xref>
            studied 18 divergent lines and 45 F
            <sub>1</sub>
            hybrids and reported that the parents VN2, BKaddi, Arka Lohit, Phule5 and LCA312 exhibited high GCA which may be utilized in recurrent selection programme for improvement in fruit quality traits.
            <xref ref-type="bibr" rid="b20">
               Rego
               <italic>et al</italic>
               . (2009)
            </xref>
            crossed eight lines of
            <italic>Capsicum baccatum</italic>
            in a complete diallel and revealed that GCA effects of the parents and SCA effects of the crosses were significant.
            <xref ref-type="bibr" rid="b12">Khalil &amp; Hatem (2014)</xref>
            evaluated six parents and 15 hybrids for yield and quality traits and found significant variance for all the traits. Estimates of GCA effects showed that the best combiner parents were found to be those of W515 and Big Dipper for fruits number, W515 and LS22 for total yield as fruits number and weight, Big Dipper for fruit diameter. For pericarp thickness and ascorbic acid content, the parental genotype Big Dipper was the best combiner, while LS22 for total soluble solids content. Estimates of SCA effects showed that the F
            <sub>1</sub>
            cross Big Dipper×B1610 reflected the highest value in all traits.
         </p>
         <p>From this prospective, the objective of the study was to estimate the general combining ability (GCA), specific combining ability (SCA) and reciprocal effects using each five quantitative and qualitative traits and to determine the most promising crosses for exploiting heterosis among the six parents.</p>
      </sec>
      <sec id="S2">
         <title>Material and methods</title>
         <p>
            The experiment was conducted in the Department of Vegetable Crops, Horticultural College and Research Institute, Tamil Nadu Agricultural University, Periyakulam, India during 2013-2015. Six homozygous inbreds
            <italic>viz</italic>
            ., Arka Lohit (P1), K1 (P2), LCA334 (P3), LCA625 (P4), PKM1 (P5) and Pusa Jwala (P6) were used as parents and the promising hybrid CO CH 1 obtained from Department of Vegetable Crops, TNAU, Coimbatore was used as standard check (
            <xref ref-type="table" rid="T1">Table 1</xref>
            ). The parents were crossed in all possible combinations, both direct and reciprocal (full diallel), to get the maximum number of hybrids. The pollen was collected from each male parent as soon as the flower buds were opened. The flower buds from plants of six female parents were emasculated in the last day evening and covered with coloured bags. In the morning, to pollinate, pollen grains from each one of the stigma of each emasculated flowers. Labels were used to identify the fruits from each different type of crosses.
         </p>
         <table-wrap id="T1">
            <label>Table 1.</label>
            <caption>
               <title>The outstanding attributes of the parents and check used for the study</title>
            </caption>
            <graphic xlink:href="sjar_e0703_t01.jpg" />
         </table-wrap>
         <p>
            The parents and progeny seeds were treated 24 hours before sowing with 4 g of
            <italic>Trichoderma asperellum</italic>
            var.
            <italic>viride</italic>
            , and sowed in 98 cells portrays. Seeds were watered with rosecan to facilitate quick germination and good growth of seedlings. The main field was prepared to a fine tilth and 25 t/ha of FYM was applied at the last ploughing. Prior to transplanting, at the time of field preparation about 2 kg/ha of
            <italic>Azospirillum</italic>
            and Phosphobacteria each were mixed with 20 kg of FYM and 30:60:30 kg/ha NPK in the form of urea, single super phosphate and muriate of potash, respectively was applied to the soil. The seedlings were transplanted in the second week of November 2013. Seedlings with age of 45 days old were transplanted with a plant spacing of 45 cm at the rate of one seedling. Fifty plants of each of the thirty F
            <sub>1</sub>
            s, six of parents were planted at a distance of 60×45 cm in a randomized block design with three replications during November 2014 to April 2015. Additionally 30 kg N/ha was given in equal splits on 30, 60 and 90 days after planting. Soil moisture was maintained during the growing season with flood irrigation at 5 days intervals. The recommended package of practices including plant protection measures obligatory to raise the good crop were followed in field (
            <xref ref-type="bibr" rid="b2">CPG,2013</xref>
            ).Data were collected from individual plants of F
            <sub>1</sub>
            generation of hot pepper for each five quantitative traits
            <italic>viz</italic>
            ., branches per plant, fruits per plant, fruit shape, individual dry pod weight, dry pod yield per plant.
         </p>
         <p>
            The observation for five quality characters were recorded from fifteen competitive plants selected randomly from each replication. The ascorbic acid content of hot pepper pod was estimated as per the standard procedure (
            <xref ref-type="bibr" rid="b25">Sadasivam &amp; Manickam, 1992</xref>
            ) and expressed in mg/100 g. The capsaicin content (%) of dry pod was estimated adopting the procedure given by
            <xref ref-type="bibr" rid="b25">Sadasivam &amp; Manickam (1992)</xref>
            .The extractable colour in hot pepper was estimated as per the procedure of
            <xref ref-type="bibr" rid="b34">Woodbury (1997)</xref>
            and expressed in ASTA units. Oleoresin content was analysed as per the procedure of
            <xref ref-type="bibr" rid="b15">Mathai (1988)</xref>
            . A drop of juice was used to record the total soluble solid (°brix) with the help of hand refractometer.
         </p>
         <sec id="S2.1">
            <title>Statistical analysis</title>
            <p>
               The statistical parameters like mean, range were calculated as per the standard methods of analysis (
               <xref ref-type="bibr" rid="b18">Panse &amp; Sukhatme, 1957</xref>
               ).
            </p>
         </sec>
         <sec id="S2.3">
            <title>Diallel analysis</title>
            <p />
            <p>
               A diallel analysis was cried out using the data of six parents and resultant thirty F
               <sub>1</sub>
               hybrids. “Null hypothesis’ was tested to show that there were no genotypic differences among the progenies. This was worked out using randomized block design. In randomized block design three sources were used
               <italic>viz</italic>
               ., blocks, genotypes and error. For blocks (b) the degree of freedom was (b-1), the degrees of freedom for genotypes was (g-1). In genotypes the results is expected as s
               <sup>2</sup>
               e (error variance) +s
               <sup>2g</sup>
               (genotypic variance). The degree of freedom for error was reported as (b-1) (g-1) and their result is expected as s
               <sup>2</sup>
               e.
            </p>
         </sec>
         <sec id="S2.4">
            <title>Griffing’s combining ability analysis</title>
            <p />
            <p>
               Estimation of general and specific combining ability was done by following
               <xref ref-type="bibr" rid="b7">Griffing (1956b)</xref>
               method I of diallel analysis, which included parents, F
               <sub>1</sub>
               s and reciprocals.
            </p>
         </sec>
         <sec id="S2.5">
            <title>General combining ability effect (GCA)</title>
            <graphic id="form1" xlink:href="sjar_e0101_form1.jpg" />
            <p>where p= number of parent; Xi.=row total of parents in the array; X.i= column total of parents in the array M, x; X..= grand total of diallel table</p>
         </sec>
         <sec id="S2.6">
            <title>Specific combining ability effects (SCA)</title>
            <graphic id="form2" xlink:href="sjar_e0101_form2.jpg" />
            <p>
               where p= number of parents; Xij= array means of F
               <sub>1</sub>
               ; Xji= array means of reciprocal F1; Xi.= row total of first parent; X.i = column total of first parent; Xj. = row total of second parent; X.j = column total of second parent; X..= grand total of diallel table.
            </p>
         </sec>
         <sec id="S2.7">
            <title>Reciprocal effects (RCA)</title>
            <graphic id="form3" xlink:href="sjar_e0101_form3.jpg" />
            <p>The variances of these effects were estimated as follows:</p>
            <graphic id="form4" xlink:href="sjar_e0101_form4.jpg" />
            <p>The square root of the variance provides the corresponding standard error for calculating critical differences.</p>
         </sec>
      </sec>
      <sec id="S3">
         <p>Results and discussion</p>
         <sec id="S3.1">
            <title>GCA effects of the parents</title>
            <p>
               The success of any plant breeding programme greatly depends on right choice of parents. The potential of a variety is judged by comparing the mean performance and combining ability of the parents (
               <xref ref-type="bibr" rid="b29">Singh, 1994</xref>
               ).
               <xref ref-type="bibr" rid="b11">Kadambavanasundaram (1980)</xref>
               suggested that the parents with high
               <italic>per se</italic>
               performance may not always be able to transmit their superior traits into hybrids and so assessment of combining ability is most needed. General combining ability (GCA) of a parent is a factor that predicts the performance of a parent over a series of cross combinations. Since GCA of a genotype is due to additive and additive×additive interaction effects, the ultimate goal of any breeder could be either to produce a variety which is homozygous for all the desired genes or to develop such parents that would produce superior hybrid. Thus combining ability assumes greater importance to assess the genetic potentialities of a genotype.
               <xref ref-type="bibr" rid="b3">Dhillon (1975)</xref>
               had also pointed out that the combining ability had given useful information on the choice of parents in form of expected performance of the hybrids and their progenies. The combining ability studies in the present investigation revealed that the parents possessed high GCA effects for different characters. Based on the GCA effects, the best parents identified as good combiners for various characters are furnished in
               <xref ref-type="table" rid="T2">Table 2.</xref>
            </p>
            <table-wrap id="T2">
               <label>Table 2.</label>
               <caption>
                  <title>Parents with high and significant GCA effect for
yield and quality traits</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t02.jpg" />
            </table-wrap>
            <p>
               At present, development of high yielding hybrids along with quality enhancement is aimed in vegetable improvement programme. Parents with high
               <italic>per se</italic>
               and significant GCA for various characters have been highlighted in
               <xref ref-type="table" rid="T3">Table 3.</xref>
               In this study some of the parents are good general combiners for yield and quality contributing characters. From this table, it is obvious that the parents ranking top in
               <italic>per se</italic>
               did not necessarily appear as top rankers when GCA effect was considered. However, for certain traits there was considerable agreement between
               <italic>per se</italic>
               and GCA of parents. Estimates of general combining ability value for the parental lines showed that the best inbreds for each characters are as follows: K1 (P2), LCA625 (P4) for branches per plant, LCA625 (P4) , K1 (P2) for fruit number, K1 (P2), PKM1 (P5) for fruit shape, K1 (P2), LCA625 (P4) for dry pod weight, LCA625 (P2), K1 (P2) for dry pod yield, LCA625 (P4), Pusa Jwala (P6)for ascorbic acid, Arka Lohit (P1), Pusa Jwala (P6) for capsaicin, Pusa Jwala (P6), LCA625 (P4) for colour value, LCA625 (P4), Arka Lohit (P1) for oleoresin and LCA625 (P4), Arka Lohit (P1) for total soluble solids. By and large, the parent LCA625 (P4) had highly significant GCA for eight characters (
               <xref ref-type="table" rid="T4">Table 4</xref>
               ). For most of the characters, its mean values also corresponded with high magnitude of its GCA. The parent LCA625 (P4) could be employed in breeding programme for overall improvement. The next best choice would be K1 (P2) which had significant GCA for five characters. This parent will be a better choice when the aim is primarily to increase the fruiting characters. Involving these parents in hybridization would result in the identification of superior hybrid combinations with favourable genes and interactions for the various traits since they are under the influence of additive genes. If a parent possessed significant GCA effects for as many traits as possible, it is well chosen for hybridization rather than parents with low GCA effects or based on mean performance. The other parent chosen for hybridization should possess favourable GCA effects for other traits, so that favourable recombinants for all the traits could be obtained. If only one trait is to be improved, both the parents should have desirable gene effects for the trait. This assumption is based on the principle that additive gene action is reflected by gene effects (
               <xref ref-type="bibr" rid="b32">Sprague &amp; Tatum, 1942</xref>
               ). Certain parents which performed well by their mean performance but failed to exhibit high GCA effects. Such low association suggests non-additive gene effects in those parents which are evidenced by higher SCA variances than GCA variances. This finding agrees with that reported by
               <xref ref-type="bibr" rid="b12">Khalil &amp; Hatem (2014).</xref>
            </p>
            <table-wrap id="T3">
               <label>Table 3.</label>
               <caption>
                  <title>Mean performance and general combining ability effects of parents for yield and quality characters</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t03.jpg" />
            </table-wrap>
            <table-wrap id="T4">
               <label>Table 4.</label>
               <caption>
                  <title>Best performing parents based on per se values and GCA effect in chilli</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t04.jpg" />
            </table-wrap>
         </sec>
         <sec id="S3.2">
            <title>Evaluation of hybrids based on specific combining ability and reciprocal effects</title>
            <p>
               The estimated results of the analysis of variance for general and specific combing abilities and reciprocal combining effects are presented in
               <xref ref-type="table" rid="T5">Table 5.</xref>
               The estimates of GCA and SCA variances help to infer the type of gene action and relative importance of the character expression in breeding programme. Magnitude of SCA variance was greater than GCA variance for all the quality traits, indicating the preponderance role of non-additive type of gene action in the expression of these traits. The results of the present study are in consonance to the findings of
               <xref ref-type="bibr" rid="b13">
                  Khereba
                  <italic>et al.</italic>
                  (2008)
               </xref>
               and
               <xref ref-type="bibr" rid="b10">Jaya Rame Gowda (2009).</xref>
            </p>
            <table-wrap id="T5">
               <label>Table 5.</label>
               <caption>
                  <title>Analysis of variance for combining ability mean
squares in chilli</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t05.jpg" />
            </table-wrap>
            <p>
               The hybrid P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               was the best performing for branches per plant on the basis of their SCA effects and mean values (
               <xref ref-type="table" rid="T7">Table 7</xref>
               ) and this hybrid was based on low×low GCA effects. The reciprocal hybrids P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>3</sub>
               ×P
               <sub>1</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               were the best performing on the basis of their RCA effect and mean values and these reciprocal hybrids were the consequential of high×high, low×low and low×low GCA effects. The
               <italic>per se</italic>
               performance was found to be high for reciprocal cross P4×P2 and this cross showed the role of additive×additive gene action. Such combinations could be used for selecting recombinants as pure lines in the later generations. Additive gene action for this trait was also reported by
               <xref ref-type="bibr" rid="b1">Asish Ghosh (2005)</xref>
               .
            </p>
            <table-wrap id="T7">
               <label>Table 7.</label>
               <caption>
                  <title>
                     <i>Per se</i>
                     performance and specific combining ability effects of direct and reciprocal crosses for yield traits
                  </title>
               </caption>
               <graphic xlink:href="sjar_e0703_t07.jpg" />
            </table-wrap>
            <p>
               The selection for high yielding genotype should be based mainly on the fruits per plant (
               <xref ref-type="bibr" rid="b6">
                  Gill
                  <italic>et al</italic>
                  ., 1973
               </xref>
               ). The hybrids P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>3</sub>
               were found to be superior based on their SCA effects and
               <italic>per se</italic>
               performance for fruits per plant and all these hybrids were the outcome of low×low GCA effects of their respective parents. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               and P
               <sub>5</sub>
               ×P
               <sub>4</sub>
               were found to be superior ones based on their SCA effects and
               <italic>per se</italic>
               performance for number of fruits per plant (
               <xref ref-type="table" rid="T7">Table 7</xref>
               ) and all these hybrids were the outcome of high×high GCA effects of their respective parents. The
               <italic>per se</italic>
               performance was found to be high for reciprocal crosses P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>5</sub>
               ×P4 and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               . All these three hybrids were good combiners with high×high parental GCA effects suggesting the influence of additive×additive gene effects on these hybrids. Hence, these parents appeared to be worthy in varietal improvement programme. It was suggested that population involving these parents, in multiple crossing programme, might be developed for isolating desirable lines. Thus, the present studies are in accordance with the outcomes of
               <xref ref-type="bibr" rid="b9">Jagadeesha &amp; Wali (2008)</xref>
               ,
               <xref ref-type="bibr" rid="b19">Prasath &amp; Ponnuswami (2008)</xref>
               and
               <xref ref-type="bibr" rid="b30">
                  Sitaresmi
                  <italic>et al</italic>
                  . (2010)
               </xref>
               .
            </p>
            <table-wrap id="T6">
               <label>Table 6.</label>
               <caption>
                  <title>Estimates of genetic components of variation for various traits in chilli</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t06.jpg" />
            </table-wrap>
            <p>
               Best performing hybrids for fruit shape based on SCA effects were P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>1</sub>
               ×P
               <sub>4</sub>
               and these hybrids were the outcome of low×low GCA effects of their corresponding parents (
               <xref ref-type="table" rid="T7">Table 7</xref>
               ). The hybrids
               <italic>viz</italic>
               ., P
               <sub>6</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>3</sub>
               ×P
               <sub>1</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               were the high performing hybrids based on RCA effects and all these hybrids were the outcome of low×high, low×low and low×high GCA effects of their respective parents. These hybrids involved one good and one poor combiner showing dominance×additive gene action which could be exploited by conventional breeding procedures such as pedigree selection.
               <xref ref-type="bibr" rid="b9">Jagadeesha &amp; Wali (2008)</xref>
               observed dominance, additive and duplicate complementary epitasis as common type of gene action for fruit shape.
            </p>
            <p>
               For individual dry pod weight, the hybrids surpassed based on the highest SCA effect were P
               <sub>2</sub>
               ×P
               <sub>6</sub>
               , P
               <sub>1</sub>
               ×P
               <sub>5</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>5</sub>
               and all these hybrids were the outcome of high×low GCA effects of their corresponding parents. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               , P
               <sub>6</sub>
               ×P
               <sub>2</sub>
               and P
               <sub>3</sub>
               ×P
               <sub>1</sub>
               were the best performing hybrids based on RCA effects and these hybrids were the outcome of low×low, low×high and low×high GCA effects of their respective parents (
               <xref ref-type="table" rid="T7">Table 7</xref>
               ). Interestingly, the
               <italic>per se</italic>
               performance was found to be high for reciprocal cross P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               which clearly indicated the significance of cytoplasmic effects controlling these trait and suggesting dominance×additive and dominance×dominance type of interaction for individual fresh fruit and individual dry pod weight. This cross could be effectively used to exploit the reciprocal differences for dry pod weight. Hence, these parents involved in the above mentioned crosses appeared to be worthy in the varietal improvement programme. It was also suggested that population involving these parents, in multiple crossing programme, might be developed for isolating desirable lines.
               <xref ref-type="bibr" rid="b19">Prasath &amp; Ponnuswami (2008)</xref>
               ; JayaRame Gowda (2009);
               <xref ref-type="bibr" rid="b30">
                  Sitaresmi
                  <italic>et al</italic>
                  . (2010)
               </xref>
               ;
               <xref ref-type="bibr" rid="b27">Savitha (2011)</xref>
               were of the opinion that the weight of fruits in chillies was governed by both additive and non additive gene effects.
            </p>
            <p>
               Dry pod yield per plant forms the major objective of the present study and the breeding procedures are focused on increasing the potentiality of this trait. The hybrids P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>3</sub>
               were found to be superior ones based on their SCA effects for dry pod yield per plant and these hybrids were the product of low×low, and high×low GCA effects of their parents. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>6</sub>
               ×P
               <sub>2</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               had the low×high GCA combination resulting in high positive significant SCA suggesting dominant×additive gene interaction. The hybrids, P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>5</sub>
               ×P
               <sub>4</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               were the best performing hybrids based on RCA effects (
               <xref ref-type="table" rid="T7">Table 7</xref>
               ) and these hybrids were the outcome of high×high, high×high and high× low GCA effects of their respective parents. The
               <italic>per se</italic>
               performance was found to be high for reciprocal cross P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , this cross showed additive×additive type of interaction. Hence, it was clearly indicated the significance of cytoplasmic effects controlling this trait and crosses could be effectively used to exploit the reciprocal differences. According to
               <xref ref-type="bibr" rid="b33">
                  Thakur
                  <italic>et al</italic>
                  . (1980)
               </xref>
               , 12 pairs of genes were responsible for governings this character. These results are in accordance with the outcomes of
               <xref ref-type="bibr" rid="b13">
                  Khereba
                  <italic>et al.</italic>
                  (2008)
               </xref>
               and
               <xref ref-type="bibr" rid="b31">
                  Somashekhar
                  <italic>et al.</italic>
                  (2008)
               </xref>
               .
            </p>
            <p>
               The high performing hybrids for ascorbic acid based on SCA effects were P
               <sub>3</sub>
               ×P
               <sub>6</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               (
               <xref ref-type="table" rid="T8">Table 8</xref>
               ) and these hybrids resulted from low×high, low×low, low×low GCA effects of their parents. The reciprocal hybrids viz., P
               <sub>5</sub>
               ×P
               <sub>4</sub>
               , P
               <sub>6</sub>
               ×P
               <sub>2</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               were the high performing hybrids based on RCA effects and these hybrids were the outcome of low× high, high×low and high×low GCA effects of their parents. The hybrid viz., P
               <sub>3</sub>
               ×P
               <sub>6</sub>
               , P
               <sub>5</sub>
               ×P
               <sub>4</sub>
               , P
               <sub>6</sub>
               ×P
               <sub>2</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               had one parent as good combiner. Here, the character is under the control of non additive gene action. The per se performance was found to be high for reciprocal cross P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               and this cross showed additive×dominance gene action, the conventional breeding technology needs some modification for capitalizing the genetic effects. In this context, instead of continuous selfing for a number of generations prior to selection, alternate intermating and selfing might be adopted to increase the span of selection. This would enhance the frequency of potential transgressive segregants in such breeding materials. These results are in corroboration with findings of
               <xref ref-type="bibr" rid="b4">Geleta &amp; Labuschagne (2006)</xref>
               and
               <xref ref-type="bibr" rid="b12">Khalil &amp; Hatem (2014).</xref>
            </p>
            <table-wrap id="T8">
               <label>Table 8.</label>
               <caption>
                  <title>
                     <i>Per se</i>
                     performance and specific combining ability effects of direct and reciprocal crosses for quality traits
                  </title>
               </caption>
               <graphic xlink:href="sjar_e0703_t08.jpg" />
            </table-wrap>
            <p>
               The best performing hybrids for capsaicin content based on SCA effects were P
               <sub>3</sub>
               ×P
               <sub>5</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>5</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>3</sub>
               and these hybrids were resultant of low×high, high×low and high×low GCA effects of their parents. The reciprocal hybrids viz., P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               , P
               <sub>6</sub>
               × P
               <sub>5</sub>
               and P
               <sub>4</sub>
               × P
               <sub>2</sub>
               were the high performing hybrids based on RCA effects and these hybrids were the outcome of high×high, high×low and low×high GCA effects of their respective parents. The combination P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               registered the highest positive SCA (
               <xref ref-type="table" rid="T8">Table 8</xref>
               ), which is a representation of high (positive) ×high (positive) GCA combination suggesting additive×additive gene interaction. This hybrid would throw desirable segregants as the additive gene effects are fixable. This was closely followed by the cross P
               <sub>3</sub>
               ×P
               <sub>5</sub>
               representing a low (negative) ×low (negative) GCA combination suggesting the action of epistasis gene action. While the cross P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>5</sub>
               and P
               <sub>2</sub>
               ×P
               <sub>3</sub>
               represents a high×low GCA combination probably by additive×dominant interaction. These hybrids had the greatest chance of producing transgressive segregants in later generation. These results were in conformity to the findings of
               <xref ref-type="bibr" rid="b19">Prasath &amp; Ponnuswami (2008)</xref>
               ,
               <xref ref-type="bibr" rid="b10">Jaya Rame Gowda (2009)</xref>
               ,
               <xref ref-type="bibr" rid="b27">Savitha (2011)</xref>
               and
               <xref ref-type="bibr" rid="b16">Munish Sharma (2012)</xref>
               .
            </p>
            <table-wrap id="T9">
               <label>Table 9.</label>
               <caption>
                  <title>Best performing F1 hybrids for fruit yield and quality traits in chilli</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t09.jpg" />
            </table-wrap>
            <p>
               In a breeding programme of hot pepper with the aim to improve the total extractable colour yield, it is not sufficient to develop genotypes with high fruit yield alone. For colour value, hybrids P
               <sub>5</sub>
               ×P
               <sub>6</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               and P
               <sub>1</sub>
               ×P
               <sub>2</sub>
               were best performing based on SCA effects and these were resultant of low×high, low×high and low × low GCA effects of their parents. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               , P
               <sub>3</sub>
               ×P
               <sub>2</sub>
               and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               were the high performing hybrids based on RCA effects and these hybrids were the outcome of high×low, low×low and high×low GCA effects of their respective parents. The
               <italic>per se</italic>
               performance was found to be high in the cross P2×P4 and showed dominant×additive gene action. For the cross P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               selection for this trait could be postponed to later generations in recombination breeding. These results are in accordance with the findings of
               <xref ref-type="bibr" rid="b19">Prasath &amp; Ponnuswami (2008)</xref>
               ;
               <xref ref-type="bibr" rid="b27">Savitha (2011)</xref>
               .
            </p>
            <p>
               The best performing hybrids based on SCA effects were P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               , P
               <sub>3</sub>
               ×P
               <sub>6</sub>
               and P
               <sub>1</sub>
               ×P
               <sub>5</sub>
               . These hybrids were the outcome of low×high, low×low and high×low GCA effects of their respective parents for oleoresin. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>5</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               were the best performing hybrids based on RCA effects and these hybrids were the outcome of low×low and high×low GCA effects of their respective parents. The
               <italic>per se</italic>
               performance were found to be high for direct cross P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               and this cross showed dominance×additive gene action and to exploit this genetic variance, conventional breeding procedures such as pedigree selection could be worthful. These results were in conformity with the findings of
               <xref ref-type="bibr" rid="b26">
                  Saritha
                  <italic>et al</italic>
                  . (2005)
               </xref>
               ,
               <xref ref-type="bibr" rid="b27">Savitha (2011)</xref>
               and
               <xref ref-type="bibr" rid="b16">Munish Sharma (2012).</xref>
            </p>
            <p>
               The hybrids P
               <sub>5</sub>
               ×P
               <sub>6</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               and P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               were found to be superior hybrids based on their SCA effects and
               <italic>per se</italic>
               performance for total soluble solids content. These hybrids were the products of low×high, low×high and high×low GCA effects of their individual parents. The reciprocal hybrids
               <italic>viz</italic>
               ., P
               <sub>3</sub>
               ×P
               <sub>1</sub>
               , P
               <sub>6</sub>
               ×P
               <sub>3</sub>
               and P
               <sub>3</sub>
               ×P
               <sub>2</sub>
               were the best performing hybrids based on RCA effects and these hybrids were the outcome of low×high, high×low and low×low GCA effects of their respective parents. The
               <italic>per se</italic>
               performance was found to be high for direct crosses P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               and P
               <sub>5</sub>
               ×P6 which showed dominance×additive gene action. Hence biparental breeding would be effective for exploitation of this trait. These hybrids could produce desirable transgressive segregants in the advanced generation. For harnessing non additive gene action in these hybrids, cyclic method of breeding involving selected recombinants and their inter crossing would be more desirable for improving total soluble solids content. Present results were in conformity with finding of
               <xref ref-type="bibr" rid="b12">Khalil &amp; Hatem (2014).</xref>
            </p>
            <table-wrap id="T1">
               <label>Table 10.</label>
               <caption>
                  <title>Promising F1 hybrids for fruits, quality and high
yield in chilli</title>
               </caption>
               <graphic xlink:href="sjar_e0703_t010.jpg" />
            </table-wrap>
            <p>
               The F
               <sub>1</sub>
               combinations that surpassed their parents for maximum number of components of merit were P
               <sub>4</sub>
               ×P
               <sub>2</sub>
               (LCA625×K1), P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               (K1×Arka Lohit), P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               (Pusa Jwala×PKM1), P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               (Arka Lohit×LCA334) and P
               <sub>2</sub>
               ×P
               <sub>4</sub>
               (K1×LCA625). Further, it was also found that the hybrids P4×P2 (LCA625×K1) and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               (Pusa Jwala×PKM1) were adjudged as the best crosses based on their number of fruits, individual dry weight and yield which are considered as the main objectives of development of hybrids with high yield and quality. The next best hybrid was P
               <sub>1</sub>
               ×P
               <sub>3</sub>
               (Arka Lohit×LCA334) as evidenced from better scores of number of fruits, fruit length, yield and quality traits. For quality improvement coupled with yield enhancement, the cross P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               (K1×Arka Lohit) was found to be best as evidenced from the scores obtained for all quality traits and yield in the present investigation (Tables 9 and 10).
            </p>
            <p>
               In conclusion, analysis of the GCA effects of parents for ten traits studied revealed that P
               <sub>4</sub>
               (LCA625), P
               <sub>2</sub>
               (K1) and P
               <sub>5</sub>
               (PKM1) were the best general combiners for almost all the traits. The hybrids P
               <sub>4</sub>
               × P
               <sub>2</sub>
               (LCA625×K1), P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               (K1×Arka Lohit) and P
               <sub>6</sub>
               ×P
               <sub>5</sub>
               (Pusa Jwala×PKM1) were found to be the best combiners for yield and its attributes. The hybrid P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               (K1×Arka Lohit) was the best reciprocal combiner for quality parameters based on their better mean performance and combing ability. Significant SCA and
               <italic>per se</italic>
               performance of hybrids P4×P
               <sub>2</sub>
               , P
               <sub>2</sub>
               ×P
               <sub>1</sub>
               and P6×P5 indicates there is an opportunity for developing F1 Hybrids. Further, isolation will enhance the genetic base of the populations by finding new recombinants to increase the yield and fruit quality
            </p>
         </sec>
      </sec>
   </body>
   <back>
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