Characterization and Agronomic Evaluation of Naturally Occurring … · 2020. 10. 11. ·...

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INTRODUCTION With more than 480,000 ha production area and 6.1 million tons total production, the Philippines ranks 6 th in the world’s top banana producers after India, China, Indonesia, Brazil, and Ecuador (FAO 2019b). In 2018, the total banana export worldwide reached 19.2 million tons, with the Philippines contributing 18% of the global shipment (FAO 2019a), catering to Asian countries like Japan, China, South Korea, the United Arab Emirates, Saudi Arabia, Iran, Iraq, and Kuwait. Among the banana cultivars in the country, the Cavendish, Saba, and Lakatan are the most popular. Cavendish cultivar is primarily grown for export to other countries. In terms of land area, Saba ranks first as it occupies Keywords: diversity, HCPC, Musa, PCA, Shannon-Weaver Lavernee S. Gueco 1 *, Irish B. Posada 1 , Frelyn V. Corpuz 1 , Lyka A. Yanos 1 , Felipe S. dela Cruz Jr. 1 , Olivia P. Damasco 1 , Michelle Lyka V. Descalsota 1 , and Agustin B. Molina Jr. 2 1 National Plant Genetic Resources Laboratory Institute of Plant Breeding, College of Agriculture and Food Science University of the Philippines Los Baños, Laguna 4031 Philippines 2 International Consultant for Banana R&D and Technical Adviser Department of Agriculture, Quezon City 1100 Philippines One of the major constraints in the Philippine banana chips industry is the lack of steady supply of raw materials for sustained production operation. The current cultivar, Saba, is tall and bears first fruits in almost two years after planting. This study was conducted to characterize the putative short-statured Saba germplasm assembled at the national repository, identify promising accessions based on agro-morphological traits, and evaluate their performance in replicated trials. Nineteen (19) Saba accessions were characterized morphologically using 17 qualitative and 17 quantitative descriptors. Characterization data revealed an intermediate genetic variation within the collection. The computed Shannon-Weaver diversity index ranged from 0.13 (pseudostem color) to 0.85 (wing type). Further analysis of the quantitative data grouped the accessions into six morphological clusters based on HCPC. Four promising lines were selected from the clusters formed, characterized by shorter plant height, shorter cropping period, and acceptable fruit and yield characters. The replicated evaluation trial of the selected accessions along with two check varieties revealed accession 09-063 as the most promising collection with the shortest plant height, earliest maturity, and with the plant crop's computed bunch weight per annum (BWPA) of 14.91 kg. Utilization of this accession in banana plantations, in combination with high-density planting, would provide a significant impact on yield, profit, and availability of raw Saba for direct consumption and processing. Characterization and Agronomic Evaluation of Naturally Occurring Short-statured Saba Banana in the Philippines *Corresponding Author: [email protected] 981 981 981 Philippine Journal of Science 149 (3-a): 981-992, October 2020 ISSN 0031 - 7683 Date Received: 17 Apr 2020

Transcript of Characterization and Agronomic Evaluation of Naturally Occurring … · 2020. 10. 11. ·...

  • INTRODUCTIONWith more than 480,000 ha production area and 6.1 million tons total production, the Philippines ranks 6th in the world’s top banana producers after India, China, Indonesia, Brazil, and Ecuador (FAO 2019b). In 2018, the total banana export worldwide reached 19.2 million

    tons, with the Philippines contributing 18% of the global shipment (FAO 2019a), catering to Asian countries like Japan, China, South Korea, the United Arab Emirates, Saudi Arabia, Iran, Iraq, and Kuwait.

    Among the banana cultivars in the country, the Cavendish, Saba, and Lakatan are the most popular. Cavendish cultivar is primarily grown for export to other countries. In terms of land area, Saba ranks first as it occupies

    Keywords: diversity, HCPC, Musa, PCA, Shannon-Weaver

    Lavernee S. Gueco1*, Irish B. Posada1, Frelyn V. Corpuz1, Lyka A. Yanos1, Felipe S. dela Cruz Jr. 1, Olivia P. Damasco1, Michelle Lyka V. Descalsota1,

    and Agustin B. Molina Jr.2

    1National Plant Genetic Resources Laboratory Institute of Plant Breeding, College of Agriculture and Food Science University of the Philippines Los Baños, Laguna 4031 Philippines2International Consultant for Banana R&D and Technical Adviser

    Department of Agriculture, Quezon City 1100 Philippines

    One of the major constraints in the Philippine banana chips industry is the lack of steady supply of raw materials for sustained production operation. The current cultivar, Saba, is tall and bears first fruits in almost two years after planting. This study was conducted to characterize the putative short-statured Saba germplasm assembled at the national repository, identify promising accessions based on agro-morphological traits, and evaluate their performance in replicated trials. Nineteen (19) Saba accessions were characterized morphologically using 17 qualitative and 17 quantitative descriptors. Characterization data revealed an intermediate genetic variation within the collection. The computed Shannon-Weaver diversity index ranged from 0.13 (pseudostem color) to 0.85 (wing type). Further analysis of the quantitative data grouped the accessions into six morphological clusters based on HCPC. Four promising lines were selected from the clusters formed, characterized by shorter plant height, shorter cropping period, and acceptable fruit and yield characters. The replicated evaluation trial of the selected accessions along with two check varieties revealed accession 09-063 as the most promising collection with the shortest plant height, earliest maturity, and with the plant crop's computed bunch weight per annum (BWPA) of 14.91 kg. Utilization of this accession in banana plantations, in combination with high-density planting, would provide a significant impact on yield, profit, and availability of raw Saba for direct consumption and processing.

    Characterization and Agronomic Evaluation of Naturally Occurring Short-statured

    Saba Banana in the Philippines

    *Corresponding Author: [email protected]

    981981981

    Philippine Journal of Science149 (3-a): 981-992, October 2020ISSN 0031 - 7683Date Received: 17 Apr 2020

  • almost 200,000 ha of land in the country. In general, most cooking banana varieties in the Philippines belong to the Saba subgroup, including Cardaba and Dippig. Cardaba is more popular in the southern part of the Philippines while Dippig is a popular farmer’s cooking-type cultivar in the Ilocos region. These two cultivars are the main source of raw materials for banana chips processing. Apart from this, Dippig and Cardaba are known for a variety of other uses in food preparation. Both are mostly eaten boiled/steamed but may also be eaten raw (uncooked) when ripe. They are commonly used in the preparation of popular Filipino snack foods like banana cue, “turon,” “maruya,” among others. Their leaves are used in food wrappings while its male bud or “puso ng saging” is the preferred variety for vegetable dishes like “kare-kare.” Lakatan, on the other hand, is the most popular dessert variety and sold at a higher price in the local market making production very profitable. Of the three major banana varieties grown in the country, fresh Cavendish and by-products of Saba contribute largely to the country’s export earnings. Among the processed fruit products, banana chips is the top dollar earner (Eusebio and Carpio 2002).

    One of the problems hampering the Philippines banana chips industry is the lack of raw materials for processing. The local cultivar Saba is tall, usually more than 4 m in height – and, in some cases, may reach 8 m – and have a long cropping period with an average harvest time of 644 days (DA 2018; Valmayor et al. 2002). Tall cultivars are also more vulnerable to damage caused by strong winds and typhoons. The Philippines has an average of 20 tropical cyclones every year causing significant damage in Saba growing areas, especially in Luzon. This results in an unsteady supply of raw materials for banana chips both in the domestic and foreign markets.

    Crop improvement in banana focuses on conventional breeding, mutation breeding, and genetic modification (Swenen et al. 2020). Genetic modification of Cavendish banana, the most popular variety for the export market, with high levels of pro-vitamin A (Dale et al. 2020a) and transgenic Cavendish banana with resistance to Fusarium Wilt TR4 are currently being evaluated (Dale et al. 2020b). The variety GN-60A, a mutant of Gran Naine, is an early flowering line that was regenerated from gamma-irradiated populations. The clone was micropropagated and evaluated in Malaysia and an early flowering line (10 weeks earlier) was selected and was registered under the name Novaria (Mak et al. 1995). Several putative mutant lines with height reduction, FOC tolerance, larger fruits size, earliness, and tolerant to toxins have been obtained using gamma rays and somaclonal variation (Roux et al. 2004). Resistance to the Banana Bunchy Top Virus in Lakatan has been generated through gamma irradiation coupled with in vitro techniques (Damasco et al. 2020).

    Dwarf or short-statured banana has also been reported in several cultivars – including Cachaco Enano, Prata Ana, and Figue Rose Naine – which all have equivalent normal stature counterparts (Engelborghs et al. 2004).

    Plants with short stature are more tolerant of strong wind damage and amenable to closer planting distance, thus increasing the number of plants that can be grown in any given area. Short-statured plants that are early maturing and high yielding are preferred planting materials, especially in areas frequently visited by typhoons. Since the conventional breeding of banana is expensive, slow, and complicated, one strategy for crop improvement is to collect the naturally occurring genetic variants, which could have been products of spontaneous mutations and/or selections for several years of cultivation, and evaluate them in the field under optimal growing condition.

    There is a wide range of variations in the Cavendish subgroup, including differences in pseudostem height, and the occurrence of dwarfism as a result of mutation is common (OGTR 2008). The diversity in the Philippine Saba has been reported by dela Cruz et al. (2020) and Dolairas-Laraño et al. (2018). In the farmer’s fields, some Saba cultivars with short stature have been observed to occur naturally. This results in earlier fruit production and tolerance to natural disasters such as typhoons. These banana cultivars may provide important materials for the banana chips industry. The present study aimed to (1) evaluate the diversity existing in the collection of short-statured Saba accessions using agro-morphological traits, (2) identify promising accessions with reduced cropping period and improved yield, and (3) evaluate the performance of the selected accessions in replicated trials under optimal environment.

    MATERIALS AND METHODS

    Plant MaterialNineteen (19) short-statured Saba accessions collected from different parts of the country and planted at the field genebank of the National Plant Genetic Resources Laboratory (NPGRL), Institute of Plant Breeding (IPB), University of the Philippines Los Baños (UPLB) were used in this study. The accession number, local name, and sampling location of the assembled germplasm are given in Table 1.

    Characterization and Estimation of DiversityShort-statured Saba accessions established at the NPGRL-IPB, UPLB (14°09'54.7"N, 121°15'05.7"E) were morphologically characterized using a subset of descriptors for Musa published by Bioversity International

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  • (IPGRI-INIBAP-CIRAD 1996). A total of 34 descriptor traits, 17 qualitative and 17 quantitative, were used to assess the level of diversity in the collection. Estimates of diversity were measured using the standardized Shannon-Weaver diversity index ( ) (Shannon and Weaver 1949) following the formula:

    where pi is the frequency proportion of each descriptor state and n is the number of states for each descriptor. Computed values were classified as high ( ), intermediate , or low , as described by Eticha et al. (2005).

    For quantitative characters, 10 frequency classes were determined using the mean (x) and standard deviation (σ) values (Pecetti et al. 1992), where the range of class 1 was defined as X1 < x – 2σ while the range of class 10 as X1 > x + 2σ. The diversity index for quantitative traits was computed following the formula above.

    To better visualize the quantitative data, the hierarchical clustering of principal components (HCPC) was carried out. For this purpose, principal component analysis (PCA) was initially performed to reduce the dimension of the data and determine the most significant principal

    components (PCs) for clustering. PCs with eigenvalues above 1 were retained for succeeding analysis (Kaiser 1958). The classification of accessions in different clusters was based on Ward’s criterion (Husson et al. 2010). The FactoMineR package (Le et al. 2008) was used to run the HCPC function and generated data were visualized with factoextra (https://CRAN.R-project.org). All processes in the multivariate analyses were performed under R 3.5.3 environments (https://www.R-project.org).

    Evaluation of Selected Accessions in Replicated TrialsMorphological characterization results were used to identify promising short-statured Saba accessions. To further evaluate the yield and yield characteristics of the selected accessions, a replicated trial was conducted at the experimental field of UPLB. For this purpose, clean and healthy suckers from the field genebank were collected and cultured in vitro following the published protocol (Damasco and Barba 1985) and optimized procedure developed by NPGRL Tissue Culture Laboratory (Damasco et al. 2012). This optimized protocol entails a shortened period between subculture until proliferation. Rooted meriplants were potted out and acclimatized in the nursery for 2–3 months prior to transplanting in the field. Plants with 4–5 leaves were laid out in a randomized complete block design with three blocks, each

    Table 1. List of characterized short-statured Saba accessions assembled at NPGRL-IPB, their local name, and place of collection.

    Accession no. Local name Collection site Latitude Longitude Elevation

    09-011 Cardaba Davao del Norte N07° 30' 33.7" E125° 36' 6" 71

    09-041 Dalian Davao City N07° 06' 12" E125° 27' 36" 335

    09-042 Abuhon/Cardaba Davao City N07° 06' 12" E125° 27' 36" 335

    09-043 Cardaba Davao City N07° 06' 12" E125° 27' 36" 335

    09-044 Cardaba Bukidnon N07° 51' 18" E125° 02' 4" 441

    09-056 Dippig Quirino N16° 21' 57" E121° 42' 26" 233

    09-062 Luyluy/Dippig Nueva Vizcaya N16° 29' 13" E121° 15' 32" 330

    09-063 Luyluy/Dippig Nueva Vizcaya N16° 29' 13" E121° 17' 32 334

    09-064 Luyluy/Dippig/Cardaba Nueva Vizcaya N16° 29' 12" E121° 17' 49" 342

    10-001 Saba Occidental Mindoro N13° 13' 13" E120° 36' 37" 32

    10-006 Saba Occidental Mindoro N12° 49' 46" E120° 55' 11" 30

    10-009 Saba Occidental Mindoro N13° 08' 54" E120° 41' 42" 6

    10-010 Saba Occidental Mindoro N13° 10' 51" E120° 40' 12" 28

    10-038 Sabang Binong Batangas N14° 05' 19" E121° 03' 52" 57

    10-040 Sabang Barako Batangas N13° 54' 29.7" E120° 51' 36.9" 13

    10-041 Sabang Pulpol Cavite N14° 11' 23.4" E120° 49' 38.3" 225

    10-043 Sabang Pulpol Cavite N14° 14' 1.3" E120° 46' 38.7" 109

    10-049 Saba Oriental Mindoro N12° 37' 48.9" E121° 30' 14" 10

    10-050 Saba Oriental Mindoro N12° 40' 25.5" E121° 30' 27.6" 40

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  • block represents the complete set of accessions and two additional tall cultivars belonging to the Saba subgroup, Cardaba and Dippig, as checks. Cardaba was the common cultivar being sold by tissue cultured laboratories in Davao while Dippig came from Quirino province. Each accession was replicated nine times in each block. Plants were established in the field at 3 m x 4 m spacing. At planting, 11 g of complete fertilizer (14-14-14) per plant was applied. Fertilization was followed every two months thereafter at a rate of 11 g complete fertilizer, 26 g of urea (46-0-0), and 23 g of muriate of potash (0-0-60). Weeding, stem and mat sanitation, deleafing, desuckering, and debudding were done as necessary to maintain the normal growth and development of the plants. Irrigation was done whenever necessary, especially during the dry season. Drainage canals were also constructed to remove excess water and prevent waterlogging in the experimental field. Data on selected agronomic (plant height, pseudostem girth, days to flowering, days to harvest) and yield parameters (bunch weight, BWPA, number of hands, number of fingers) were recorded during harvest.

    Analysis of variance was performed using the R 3.5.3 software (https://www.R-project.org) and significant differences between accessions were evaluated with the least significant difference set at α = 0.05.

    RESULTS AND DISCUSSION

    Morphological Diversity of Short-statured Saba GermplasmThe computed diversity indices for quantitative traits ranged from 0.31 for fruit length to 0.67 for bunch weight and total soluble solids, with a mean diversity value of 0.54. In addition to fruit length, the quantitative traits “days from shooting to harvest” and “days from planting to harvest” also showed low diversity values of 0.34 and 0.38, respectively. High level of diversity on the other hand was also observed for pseudostem height (0.61), pseudostem girth (0.65), fruit width (0.63), fruit peel weight (0.65), fruit peel thickness (0.64), and pedicel length (0.61). The remaining quantitative characters were classified as intermediate, with diversity values ranging from 0.41–0.58. The overall diversity mean of 0.54 implies an intermediate variation within the selected short-statured Saba in terms of quantitative characters (Table 2a). Qualitative traits were observed to have low (eight characters) to intermediate (six characters) diversity, with values ranging from 0.13–0.50. Only three characters, i.e. petiole canal leaf, wing type, and bunch appearance, showed high variation with H’ values of 0.64, 0.85, and 0.66, respectively (Table 2b). The diversity in the bunch appearance of the different accessions is shown in Figure 1. Low diversity index values are indicative of the dominance of a particular descriptor state over the others, while high diversity index values indicate a relatively even distribution of different descriptor states. The dominant

    Table 2a. Diversity of quantitative traits among short-statured Saba accessions, as revealed by 𝐻′.Quantitative trait Mean Range H' Diversity level

    Pseudostem height (m) 3.4 2.76–3.95 0.61 High

    Pseudostem girth (cm) 68.1 49–80 0.65 High

    Fruit weight (g) 136.2 89.3–237.9 0.58 Intermediate

    Fruit length (mm) 66.1 10.5–163.4 0.31 Low

    Fruit width (mm) 46.8 42.2–51.5 0.63 High

    Fruit thickness (mm) 37.9 33.0–44.8 0.53 Intermediate

    Fruit peel weight (g) 50.5 30.6–74.6 0.65 High

    Fruit peel thickness (mm) 2.9 1.8–4.2 0.64 High

    Pedicel length (mm) 26.7 22.1–30.8 0.61 High

    Pedicel width (mm) 12.2 9.3–13.6 0.61 High

    Total soluble solids (°Bx) 25.9 22.3–30 0.67 High

    Bunch weight (kg) 20.2 11.6–29.2 0.67 High

    Hand number 8.6 6.0–10.0 0.45 Intermediate

    Finger number 143.6 62–184 0.41 Intermediate

    Days from planting to shooting 286.4 198–333 0.46 Intermediate

    Days from shooting to harvest 140.7 82–256 0.34 Low

    Days from planting to harvest 427.1 351–454 0.38 Low

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  • Table 2b. Diversity of qualitative traits among short-statured Saba accessions, as revealed by 𝐻′.

    Qualitative trait Number of descriptor states

    Dominant character H' Diversity level

    Pseudostem color 16 Medium green 0.13 Low

    Pseudostem appearance 2 Shiny 0.31 Low

    Pseudostem underlying color 8 Watery green 0.17 Low

    Underlying pigmentation 8 Pink purple 0.38 Low

    Blotches at petiole base 5 Small blotches 0.22 Low

    Petiole canal leaf 5 Margins curved inward 0.64 High

    Petiole margins 5 Winged and clasping 0.41 Intermediate

    Wing type 2 Dry 0.85 High

    Edge of petiole margin 2 Colorless 0.50 Intermediate

    Appearance of leaf lower surface 2 Dull 0.50 Intermediate

    Wax on leaves 4 Moderately waxy 0.15 Low

    Shape of leaf blade base 3 Both sides rounded 0.32 Low

    Bunch position 5 Slightly angled 0.41 Intermediate

    Transverse section of fruit 3 Pronounced ridges 0.48 Intermediate

    Fruit apex 5 Blunt-tipped 0.45 Intermediate

    Bunch shape 5 Truncated cone shape 0.37 Low

    Bunch appearance 3 Lax 0.66 High

    Figure 1. Diversity in bunch appearance of select accessions: (L–R) accessions 09-063, 09-041, and 10-050.

    character for each qualitative trait measured is shown in Table 2b.

    Sustainable conservation and effective use of germplasm collections for research and breeding necessitate prior knowledge of the diversity existing in the Genebank (Beuselinck and Steiner 1992). Hence, characterization, evaluation, and classification are crucial components for the management of genetic resources (Ortiz 1997).

    Morphological characterization is a simple way to evaluate variability in germplasm collections (Khan et al. 2015) and has been widely used in estimating the degree of diversity in different plant species (Archak et al. 2016; Santos et al. 2012; Upadhyaya 2003; Wei et al. 2019; Zavinon et al. 2019). Understanding the genetic relatedness among individuals facilitates the selection of superior materials for direct utilization or as parents in various breeding programs.

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  • Multivariate Analysis of Quantitative TraitsHCPC was performed to describe the existing variability in the short-statured Saba collection and elucidate the similarities between individuals based on 17 quantitative characters. Eigenvalues above 1 were observed in the first four PCs (Table 3), explaining 82.2% of the total variability. The first PC, which explains 34.6% of the total variance, is positively correlated with pseudostem height (PSEHT), pseudostem girth (PSEGRT), fruit peel thickness (PELTH), total soluble solids (TSS), hand number (HNDNO), finger number (FINNO), days from planting to shooting (DPTOSH), and days from planting to harvest (DPTOH). On the other hand, fruit weight (FRTWT), fruit length (FRTLN), fruit thickness (FRTTH), and fruit peel weight (PELWT) were found negatively correlated to PC1. The second principal component accounted for 21.3% of the variance and was positively correlated with plant height, fruit width (FRTWD), fruit thickness, fruit peel weight, fruit peel thickness, pedicel length (PEDLN), and pedicel width (PEDWD). PC3 was mostly positively correlated with yield characters including bunch weight (BUNWT), hand number, and finger number. For PC4, only total soluble solids and days from shooting to harvest (DSHTOH) were associated at

    –0.59 and –0.74, respectively. Figure 2 presents the PCA graph of the variables and individuals plotted on the first two components. From the graph, it can be seen that PC 1 opposes individuals with high values of PELTH, DPTOSH, PSEGRT, and DPTOH (e.g. 10-006, 10-009) to individuals with high values of FRTWT, FRTTH, PELWT, and FRTLN (e.g. 10-040 and 10-038), with the former characterized by a strongly positive coordinate to the right of the axis and the latter categorized with strongly negative coordinate to the left of the axis. Conversely, PC2 opposes individuals with low values for the variable FRTLN to individuals, with low values of PEDWD, PELTH, PEDLN, and PELWT placing individuals from the latter at the top of the graph (positive coordinate) and the former at the bottom (negative coordinate).

    Succeeding cluster analysis was performed in the first four PCs and the classification made on the individuals revealed six morphological clusters (Figure 3). Among the six clusters formed, the 1st and 3rd clusters were observed to include a single accession, i.e. 10-038 and 09-063, respectively. Conversely, clusters 5 and 6 were found to encompass 42% and 26% of the total accessions evaluated, respectively. The lone accession in cluster 1

    Table 3. Correlation coefficients between each analyzed trait and the first four PCs, along with eigenvalues and proportion of variability explained by each PC.

    Trait Trait code PC1 PC2 PC3 PC4

    Pseudostem height PSEHT 0.46 0.58 0.31 0.05

    Pseudostem girth PSEGRT 0.84 0.26 0.16 0.31

    Fruit weight FRTWT –0.85 0.39 0.14 0.27

    Fruit length FRTLN –0.83 –0.36 0.27 0.05

    Fruit width FRTWD –0.28 0.75 0.28 0.10

    Fruit thickness FRTTH –0.78 0.52 0.11 0.03

    Fruit peel weight PELWT –0.62 0.61 0.02 0.41

    Fruit peel thickness PELTH 0.60 0.47 –0.40 0.11

    Pedicel length PEDLN 0.05 0.61 0.09 –0.29

    Pedicel width PEDWD 0.12 0.76 –0.02 0.07

    Total soluble solids TSS 0.47 0.31 0.30 –0.59

    Bunch weight BUNWT 0.06 0.23 0.90 0.07

    Hand number HNDNO 0.66 –0.10 0.69 –0.07

    Finger number FINNO 0.59 –0.23 0.72 0.04

    Days from planting to shooting DPTOSH 0.77 0.09 –0.40 0.42

    Days from shooting to harvest DSHTOH –0.38 0.39 –0.19 –0.74

    Days from planting to harvest DPTOH 0.60 0.43 –0.62 –0.12

    Eigenvalue 5.88 3.62 2.92 1.55

    Variability % 34.6 21.3 17.20 9.14

    Accumulated variability % 34.6 55.9 73.10 82.24

    *Values in bold indicate significant correlations at α = 0.05.

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  • is characterized by longer days from shooting to harvest, short pseudostem girth, low number of fruit hands, and fewer fruit fingers. Accessions in the 2nd cluster were found to have heavier individual fingers, high values for fruit peel weight, and low total soluble solids; and share the same characteristics as cluster 1 in terms of the number of hands and fingers. The accession in cluster 3 takes shorter days to harvest. Similarly, accessions in cluster 4 were characterized by shorter days to harvest, shorter days to shooting, and relatively low values for peel thickness. Accessions in cluster 5 were observed to have longer days to shooting, low values for fruit-related traits (including fruit thickness, fruit weight, and fruit

    diameter), and intermediate value for bunch weight. Accessions in the last cluster have bigger pseudostem girth and high total soluble solids. The identification of these six morphological clusters provided information on the differences and similarities among the short-statured accessions in terms of quantitative traits. As most of the traits measured presents some degree of importance for utilization and breeding, the information generated can guide researchers/breeders to select accessions for further exploitation in downstream programs.

    In terms of morphological characters, three distinct subgroups were observed. The Bluggoe types: composed of 10-038 (cluster 1), 10-040 (cluster 2), and 10-050

    Figure 2. PCA graph of variables and individuals plotted on the first two PCs. Dim1 and Dim2 correspond to PC1 and PC2, respectively.

    Figure 3. Cluster plots showing the number of clusters based on 17 quantitative traits measured.

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  • (cluster 2) have short pseudostem heights with dark green fruits but poor eating quality, locally known as Gubao/Datu types. The other group is an intermediate between the Bluggoe and Saba types, which have intermediate heights, but bunch and fruit characteristics comparable with the typical Saba/Cardaba types. Accessions belonging to this group include 09-063 (cluster 3), 09-062 (cluster 4), and 09-064 (cluster 4); the rest of the accessions may be the typical Saba/Cardaba fruit types.

    The preliminary agronomic performance of the different short-statured Saba is presented in Table 4. The accession 09-063 and 09-064 obtained the heaviest bunch weights with 29.2 kg and 26.3 kg, respectively. These two accessions likewise displayed short pseudostem heights and short crop cycle. Although some accessions like 10-038 and 10-040 have short stature, they do not have good eating quality. Another interesting accession is 10-009 with a bunch weight of 26.2 kg and with intermediate height (3.65 m). There are other accessions that are shorter, but they are Gubao types (10-038 and 10-040) and tastes are not acceptable. According to the manufacturer of banana chips, 09-044 is excellent for banana chips; hence, it was included in the replicated trial.

    Compared to the typical Saba, the selections have a shorter crop cycle (351–415 days) and shorter pseudostem height (3.04–3.65 m). Valmayor et al. (2002) reported that the pseudostem height of typical Saba measures 4.5–5.0 m tall and takes roughly 644 days to harvest. In the farmer’s field, some Saba plants were observed to reach 8–9 m. Shortened crop period and shorter pseudostem height are two of the most desirable agronomic traits in banana. The 27–32% drop in plant height and 36–45% reduction in the crop cycle would have a significant impact on the yield, profitability, and availability of raw Saba for consumption or processing. The shortened period of production equates to an earlier return on investment (ROI) and cuts a significant amount of operating cost for maintenance. Shorter plants on the other hand has lesser vulnerability to wind damage caused by typhoons and facilitates easy harvesting. Moreover, shorter plants are ideal on steep slopes and can be densely planted, creating high yield potential and economic profit (dela Cruz et al. 2009).

    Agronomic and Yield Performance of Promising AccessionsThe replicated trial revealed significant variations in the agronomic and yield performance of the different accessions (Table 5). The shortest mean height was recorded

    Table 4. Preliminary agronomic performance of different short-statured Saba accessions.

    Accession number

    Local name Pseudostemheight (m)

    Days from planting to harvesting

    Bunch weight(kg)

    Bunch weightper annum

    (kg yr–1)

    No. of hands

    No. of fingers

    09-011 Cardaba 3.46 443 23 18.95 9 142

    09-041 Dalian 3.40 454 18.1 14.55 9 147

    09-042 Abuhon/Cardaba 3.50 443 25 20.60 10 161

    09-043 Cardaba 3.33 454 21.2 17.04 9 165

    09-044 Cardaba 3.32 415 23 20.23 8 146

    09-056 Dippig 3.12 454 11.6 9.33 8 116

    09-062 Luyluy/Dippig 2.96 364 21.7 21.76 10 180

    09-063 Luyluy/Dippig 3.24 351 29.2 30.36 10 182

    09-064 Luyluy/Dippig/Cardaba 3.04 372 26.3 25.81 9 180

    10-001 Saba 3.52 454 20 16.08 9 167

    10-006 Saba 3.95 454 23 18.49 9 156

    10-009 Saba 3.65 443 26.2 21.59 9 152

    10-010 Saba 3.68 454 22.2 17.85 9 145

    10-038 Sabang Binong 2.77 454 13.2 10.61 8 138

    10-040 Sabang Barako 2.76 351 20 20.80 6 72

    10-041 Sabang Pulpol 3.51 415 16.1 14.16 9 154

    10-043 Sabang Pulpol 3.35 443 16 13.18 9 152

    10-049 Saba 3.41 454 14.2 11.42 8 130

    10-050 Saba 3.21 443 13 10.71 7 117

    *All agronomic traits measured were obtained on the main crop at harvest.

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  • Table 5. Mean values of agronomic and yield parameters of the selected promising accessions in replicated trial (n = 9).

    Parameters Cardaba Dippig 09-063 09-064 10-009 09-044

    Height (m) 4.29 bc 4.50 c 3.13 a 4.34 bc 4.14 bc 3.92 b

    Girth (cm) 73.91 a 78.46 a 59.42 b 75.24 a 76.01 a 75.53 a

    Days to shooting 409.94 b 385.90 b 286.14 a 425.31 b 383.71 b 398.78 b

    Shooting to harvest 152.71 bc 155.07 bc 118.93 a 142.09 b 159.43 c 147.89 bc

    Days to harvest 556.90 b 540.97 b 405.08 a 567.40 b 543.14 b 539.56 b

    Bunch weight (kg) 20.79 bc 26.42 a 16.55 d 19.74 c 22.80 b 20.03 bc

    Number of hands 8.58 bc 9.33 ab 10.18 a 7.82 c 8.73 bc 9.20 ab

    Number of fingers 133.04 b 157.32 ab 178.65 a 126.76 b 140.80 b 155.40 ab

    Bunch weight per annum (kg) 13.63 bc 17.83 a 14.91 bc 12.70 c 15.32 b 13.55 bc

    *Means with the same letter in a row are not significantly different at α = 0.05. All measurements were taken from the main crop.

    in accession 09-063 (3.13 m), which was significantly shorter than the check cultivars. This was followed by accession 09-044 (3.92 m). On the other hand, plant heights of cultivars 09-064 and 10-009 were 4.34 m and 4.14 m, respectively, and were comparable to Cardaba (4.29 m). Dippig exhibited the tallest pseudostem height with 4.5 m. Accession 09-063 had a height reduction of 26.23% and 30.34% as compared to the check cultivars Cardaba and Dippig, respectively. In contrast to tall banana cultivars, short-statured accessions are amenable to closer planting distance (dela Cruz et al. 2009); hence, a greater number of plants can be planted per unit area. In the cultivar Grand Nain, high-density planting resulted in an improved yield with increased bunch weight and improved cost-benefit ratio in the production cycle (Mahmoud 2013).

    The earliest onset of shooting was observed in accession 09-063 with an average of 286 days or 9.5 months and was found statistically different from all other accessions. In the check cultivars Cardaba and Dippig, the average number of days to shoot were 409.94 and 385.90 days, respectively. A similar trend was observed in terms of the number of days from planting to harvest, other cultivars are statistically the same except for 09-063. The average days to harvest were as follows (earliest to latest): 09-063 (405 days), 09-044 (540 days), Dippig (541 days), 10-009 (543 days), Cardaba (557 days), and 09-064 (567 days).

    Accession 09-063 (119 days) also had the shortest days from shooting to harvest. It was followed by 09-064 (142 days) while the rest of the evaluated accessions displayed 148–159 days from shooting to harvest. Significant differences were also found in the yield parameters gathered such as bunch weight, number of hands, and number of fingers as well as computed BWPA. In terms of the bunch weight of the plant crop, Dippig yields the heaviest with 26.42 kg. It was followed by 10-009 (20.80 kg), Cardaba (20.79 kg), and 09-044 (20.03 kg) which is comparable to each other. Accession 09-064 (19.74 kg)

    is also comparable to Cardaba while 09-063 significantly yields the least with 16.55 kg average bunch weight. Although characterized by the least average bunch weight, accession 09-063 has considerably the greatest number of hands and fingers among other cultivars with average values of 10 hands and 157 fingers. The other selections produced statistically the same number of hands and fingers compared to the check cultivar, Cardaba.

    Considering both yield and earliness to harvest of the plant crop, the BWPA [bunch wt x (365/days to harvest)] was computed. The heaviest BWPA was recorded in Dippig (17.83 kg yr-1). This was followed by 10-009 (15.32 kg yr-1), 09-063 (14.91 kg yr-1), and 09-044 (13.55 kg yr-1) which are found comparable to Cardaba (13.63 kg yr-1). The least BWPA was recorded to 09-064 with 12.70 kg yr-1. The computation of BWPA implies that 09-063, though has the lowest bunch weight per harvest, can compensate for the harvest of the typical Saba/Cardaba due to its early maturing characteristic. The accession 09-063 was not significantly different from the check cultivar Cardaba in terms of BWPA. Translating the computed BWPA into yield per hectare (14.91 * 1111 plants ha-1 = 16.56 tonnes ha-1) show an increase in the average yield of Saba by 2.56 tonnes compared to the baseline productivity of 14 tonnes ha-1 (DA 2018). But regardless of height, Dippig yields the highest bunch weight per harvest and BWPA. Dippig and 10-009 may be recommended to farmers who wanted large bunches and bigger fruits not considering the disadvantage of planting tall and late-maturing varieties (i.e. wider planting distance, susceptibility to wind damage). However, planting early maturing cultivars have the advantage of having less time exposed to pest and diseases, as well as abiotic stresses (i.e. typhoons, droughts, floods), which are expected to occur more frequently in the future due to climate change.

    Breeding banana with improved traits is difficult through conventional methods, as most cultivars are known to

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  • have high male and female sterility (Tripathi et al. 2007). In the Philippines, conventional breeding programs in banana are lacking and development of individuals with desirable agronomic traits is often carried out through induced mutations (Damasco et al. 2020; Jain 2010; Lamo et al. 2017). Aside from induced mutation, natural mutations also occur in the field, which can result in individuals harboring traits of interest. For example, the widely distributed Dwarf Cavendish (Ploetz et al. 2007) is reported to have been derived from a mutation from tall members of the Cavendish group (Constantine and Rossel 1999). Several other dwarf cultivars were described to have originated from natural mutation including Nanicão (derived from Nanica) and Prata-Anã (a mutation from Branca) (Donadio et al. 2019), whereas Nanicão Corupá and Prata-Catarina were clones from a selected natural variation in the field. The Pisang Awak subgroup (also called Kluai Namwa in Thailand and Lagkitan/Katali locally) also have a wide range of diversity ranging from Dwarf to Tall cultivars. As it is, an intensive evaluation of banana germplasm provides a simple and direct approach in identifying promising accessions/clones for cultivation.

    SUMMARY AND CONCLUSIONA total of 19 Saba germplasm collections were characterized and evaluated to identify and select accessions with short stature, reduced cropping period, and acceptable yield. Characterization data revealed an intermediate level of variation within the germplasm collection. The wide variability in the quantitative traits measured enabled the classification of the 19 accessions into six morphological clusters through HCPC. Preliminary evaluation allowed the selection of four promising accessions in terms of pseudostem height, fruit characters, cropping period, and yield. Further evaluation of these promising accessions in replicated trials revealed 09-063 as superior over the others as characterized by the shortest pseudostem height, shortest cropping period, and intermediate bunch weight per year. The identification of short-statured Saba variety is valuable, particularly in areas prone to tropical cyclones, since shorter plants are less vulnerable from damage to strong winds. In addition, short-statured plants can be densely planted, resulting in an increase in the number of plants per unit area. A shorter cropping period also results in an earlier ROI from the plant crop. Utilization of short-statured and early fruiting variety in banana plantations, in combination with high-density planting, would have a significant impact on yield, profit, and availability of raw Saba for direct consumption and processing. Still, there is a need to assess the performance of 09-063 along with other promising accessions and checks in a multi-location trial as quantitative traits are highly affected

    by different growing conditions. Finally, this study had shown that clonal selection of naturally occurring genetic variants from the farmer’s field presents a simple and direct approach for the identification of improved banana varieties. This method can be used as an alternative strategy for crop improvement.

    ACKNOWLEDGMENTSThe authors would like to thank the Department of Science and Technology – Philippine Council for Agriculture, Aquatic and Natural Resources Research and Development for funding this research. Members of the administrative staff – Paulo Jerome M. Lopez, Maria Adisaz, Melanie Bacor, John Mark Barios, Aaron Javier, Marlon Borja, Joel Evangelista, and Ceferino Evangelista (retired) – are also acknowledged for various assistance in conducting this research.

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