Phenotypic Characterization, Evaluation, and Classification of Cassava (<i>Manihot esculenta</i> Crantz) Accessions in Ethiopia

Bilate Daemo, Berhanu; Belew Yohannes, Derbew; Mulualem Beyene, Tewodros; Abtew, Wosene Gebreselassie

doi:https://doi.org/10.1155/2023/1559070

International Journal of Agronomy

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Abstract Introduction Materials and Methods Results and Discussion Conclusion Data Availability Conflicts of Interest Authors’ Contributions Acknowledgments Supplementary Materials References Copyright Related Articles

Research Article | Open Access

Volume 2023 | Article ID 1559070 | https://doi.org/10.1155/2023/1559070

Phenotypic Characterization, Evaluation, and Classification of Cassava (Manihot esculenta Crantz) Accessions in Ethiopia

Berhanu Bilate Daemo,^1,2Derbew Belew Yohannes,²Tewodros Mulualem Beyene,³and Wosene Gebreselassie Abtew²

Academic Editor: Maria Serrano

Received11 Mar 2023

Revised22 Apr 2023

Accepted22 Aug 2023

Published28 Aug 2023

Abstract

Cassava has a crucial role in benefiting smallholder farmers as the main food and income source in southern Ethiopia. Characterization accessions are crucial for assessing variation, classifying, and identifying desirable accessions for crop improvement and conservation. In this regard, there needs to be more information on the morphological characterization and classification cassava accessions. Thus, the aim of this research was to systematically characterize, evaluate, and classify cassava accessions using qualitative characters to provide useful information for breeding program and conservation. A total of 64 accessions were planted using a simple lattice design during the 2020-2021 cropping season. Thirty qualitative data were collected at 3, 6, 9, and 18 months after planting and analyzed using the SAS and R-software packages. The high variable characteristics were the shape of the central leaflet, petiole color, leaf retention, branching habit, the color of the stem epidermis, the color of the stem exterior, the external color of the storage root, and the color of the root pulp towards frequency distribution analysis. The Shannon–Weaver diversity index ranged from 0.24 to 1.47, with an overall mean of 0.84. The first three dimensions in the multiple correspondent method explained approximately 39.39% of the total variation, with Dim 1 accounting for 20.77% and Dim 2 accounting for 9.98%, while petiole color and texture of the root epidermis were the leading contributors to the total variation, respectively. In clustering analysis, 64 accessions were classified into 4 clusters of varying sizes. The distribution of accessions in each cluster revealed that 52 accessions were in cluster I, 6 in cluster II, 5 in cluster III, and 1 in cluster IV. Each cluster was varied by a major group characteristic that it represented. Furthermore, the study identified the desirable accessions for desired storage root characteristics such as root constrictions, the external color of the storage root, the color of root pulp, the color of the root cortex, and cortex peeling tendency. In conclusion, the various analyses performed indicated the existence of sufficient genetic variability for the characteristics evaluated, which could be attributed to the dissimilar genetic backgrounds of the evaluated accessions. Thus, these could be utilized for breeding work and conservation.

1. Introduction

Cassava (Manihot esculenta Crantz) grows more frequently in subtropical and tropical regions despite being native to Latin America, and it is the fourth major food crop in the world, next to maize, rice, and wheat [1]. It is regarded as an important nutrient source for ensuring food security in developing countries and a key source of food calories for two out of every five people in Africa [2]. A number of industrial products are made with cassava starch, including paper, cardboard, textiles, plywood, glue, alcohol, animal feed, nongrain starch, ethanol, and biofuel [3]. Cassava has been shown to adapt and grow well in different agroecologies in Ethiopia, with various levels of yield [4]. It is an essential food crop that provides a large portion of people’s daily consumption as well as the principal supply of carbohydrates in southern Ethiopia [5].

Cassava is a major storage root crop with a lot of potentially useful genetic variation [6]. Jahufer and Gawler [7] stated that genetic variation is central to plant breeding, as good management of variation can produce permanent gains in the yield of the plant and can buffer seasonal fluctuations. Farmers continue to keep a diverse set of accessions (landraces) on their farms, despite the fact that they may be low yielding and susceptible to some biotic and abiotic stresses [8]. These landraces may have greater genetic variability, which may enhance gene flow via hybridization [8, 9]. Thus, landraces are valuable genetic resources for breeding and other crop enhancement efforts [9, 10]. Furthermore, maintaining genetic resources is crucial for achieving genetic progress in breeding programs throughout the recombination and selection cycles [11]. To make the best use of available genetic variability, the genetic material must be properly organized and analyzed, increasing the likelihood of selecting genotypes with superior performance on the traits of interest [12]. Both morphological and agronomic variables, as well as molecular analysis, can be used to evaluate genetic variability [12]. The primary goal of characterization is to get rid of duplicate accessions, generate a genetically unique core sample for use in breeding programs, and produce progeny who have a higher level of heterotic [13].

The morphological characteristics of cassava are prominently variable, signifying a high degree of interspecific hybridization [14, 15]. Information on the genetic relationship between accessions is an important component of plant breeding programs because it provides knowledge of the genetic diversity available for producing new allelic combinations in a segregated population [11]. In cassava, it is supposed that a broad range of genetic variability was produced through centuries of farmer management [10]. Morphological characterization and evaluation of locally available accessions are indispensable for making information available for cassava root yield improvement. According to Fukuda and Guevara [16], assessing existing genetic variability is necessary and should be based on appropriate and recognised descriptors. So far, studies using qualitative characteristics conducted in various countries around the world have revealed significant variability within farmer-owned cassava cultivars [17]. According to Asare et al. [18], morphological characters are commonly used in primary evaluation because they are a quick and easy way to determine the degree of variability. These morphological characters reveal the true variability as perceived by farmers, and morphological characterization has been used to identify genetic diversity among cassava varieties [19].

Cassava is an essential food crop in Ethiopia that provides food security and income as well as a significant percentage of the daily diet [5, 20]. However, there are various research gaps in this genetic research on important morphological characteristics in cassava, which lagged far behind other root and tuber crops [5]. Such research gaps are attributed to major problems such as a lack of desired features for the cassava storage root and high-root-yielding cultivars. This highlighted the significance of extending research efforts to look at the morphological characterization and relationships among cassava accessions through the collection of available genetic resources. Therefore, the aim of this research was to characterize, evaluate, and classify cassava accessions obtained from different sources using qualitative characteristics to provide valuable knowledge for cassava improvement programs and conservation efforts.

2. Materials and Methods

2.1. The Study Area’s Description

The experiment was carried out during the 2020-2021 cropping season at the Tarcha research site in the Dawuro zone, Southwest Ethiopia People’ Regional State, and Bonbe research site in the Wolaita zone, Southern Nations Nationalities and Peoples’ Region State. Tarcha research site is situated at an altitude of 1250 meters above sea level at latitude 07°09′32″N and longitude 037° 10′16″E [21]. The average annual rainfall in the area is 1392 mm, with a mean maximum and minimum temperature of 30°C and 17°C, respectively [21]. The soil in the study area is nitosol, which is weathered brown and has a pH of 5.6. Bonbe research site is located at an altitude of 1701 meters above sea level at latitude 07°08′15.5″N and longitude 037°34′54.1″E [21]. The average annual rainfall in the area is 1450 mm, with a mean maximum and minimum temperature of 26°C and 15°C, respectively [21]. The soil in the study area is nitosol, which is weathered red and has a pH of 4.25.

2.2. Plant Materials and Design

Sixty-four cassava accessions were used for the study, fifteen were from the Nigeria and forty-nine accessions were from the Jimma and Hawassa Agricultural Research Centers (Table 1). The experiment was designed as an 8 × 8 simple lattice. Mature cassava cuttings measuring 25–30 cm long were planted in an experimental plot (7 m²) at an inter-row spacing of 1 m and an intrarow spacing of 1 m on the top of the ridge at a slanting (an angle of 45°) position. The evaluation was conducted on three plants for each accession per plot. All cultural practices were performed as recommended by Markos et al. [22] and farmers’ practices in the area.

2.3. Data Collection

The morphological characters were observed at four periods, 3, 6, 9, and 18 months after planting (MAP), based on 30 descriptors of cassava as developed by Fukuda et al. [23]. The most frequently observed variant was noted in three plants (Table 2). The presence or absence of seeds and fruit was recorded throughout the vegetative stage until harvest.

2.4. Data Analysis

Morphological frequency distributions were estimated based on the characters observed in the accessions. To assess the phenotypic diversity of all accessions, the Shannon–Weaver diversity index (H′) was computed using the phenotypic frequencies. The Shannon–Weaver was estimated using the following formula [24]:where pi is the proportion of accessions in the i^th class of the trait to the total number of accessions grouped in the trait, where “n” is the number of phenotypic classes in the trait. Multiple correspondent analyses (MCA) were performed using the method developed by Abdi and Valentin [25] to analyze a set of observations for a set of categorical variables using statistics [26]. The MCA is a method helpful for understanding the similarities between the categories of variables and the associations between the variables [27]. It displays and allows the identification of factorial axes that reveal the most discriminant variables [28]. Clustering analysis was performed based on unweighted pair group methods with an arithmetic average (UPGMA). The numbers of clusters were determined using the cubic clustering criteria (CCC) as described by Mohammadi and Prasanna [29]. The appropriate numbers of clusters were determined from the values of pseudo-F and pseudo-T² using the SAS software [26] and R–software package [30].

3. Results and Discussion

3.1. Distribution in Characters

Accessions showed high to low variations in the qualitative characters’ frequency distribution among 64 accessions (Table 3). Apical leaf color 49% of accessions were observed as purple-green, 45% showed purple, and 6% showed dark green (Table 3). Similarly, the distribution of pubescence on apical leaves was observed to be present (53.12%) and absent (46.88%) among accessions. The central leaflet shape of the accessions varied more, with 39% having a lanceolate central leaflet shape, 26% having an elliptic-lanceolate shape, 22% having an oblong-lanceolate shape, 5% having an ovoid shape, 5% being straight or linear, and 3% having an obovate-lanceolate shape (Table 3).

Similarly, the characters showed higher variability among accessions for petiole color (36% red, 25% reddish green, 22% yellowish green, 9% purple, and 8% greenish red) and leaf retention (45.31% better than average retention, 17.19% outstanding leaf retention, 17.19% both less than average and average leaf retention, and 9.38% very poor retention) (Table 3). Some characters showed medium variability, such as leaf lobe number (55% seven lobes, 39% nine lobes, and 6% five lobes), leaf vein color (75% green, 12.5% reddish green in less than half of the lobe, and 12.5% reddish green in more than half of the lobe), and petiole orientation (62.5% horizontal, 31.25% inclined upwards, and 6.25% inclined downward), while others showed low variability, such as leaf color (54.69% dark green and 45.31% light green) and lobe margins (51.56% winding and 48.44% smooth) (Table 3). Furthermore, accessions displayed wide variation in the following stem characteristics: the stem exterior color (64% silver, 28% greeny-yellowish, 5% golden, and 3% orange), the stem epidermis color (61% orange, 25% cream, and 14% light brown), the branching habit (12.5% erect, 29.69% dichotomous, 32.81% trichotomous, and 25% tetrachotomous), and the measurement between leaf scars (45.31% short, 40.63% medium, and 14.06% long) (Table 3). The stem cortex color was 95.31% light green and 4.69% orange, while the growth habits of the stem were straight (76.56%) and zigzag (23.44%).

The root shape of the accessions varied from conical-cylindrical (42%) to irregular (14%), while the root constriction showed few to none (94%) and some (6%) accessions (Table 3). White or cream (34% of the total) was the most common external color of storage roots, followed by yellow (33%), light brown (28%), and dark brown (5%), with dominant white and cream root pulp colors (Table 3). Similarly, the root cortex color was observed to be pink (44%), white or cream (31%), and yellow (25%), whereas the cortex peeling extent and texture of the root epidermis were dominantly difficult and smooth, respectively (Table 3). Cassava has few to no root constriction, white or cream external roots, yellow root pulp color, white or cream root cortex, and ease of removing the root cortex, as reported by Gomes [31], Vieira et al. [32], and Tiago et al. [33], which are desirable features for the cassava storage root from both a genetic improvement and an agronomic point of view. According to Table 3, our study found that 94% (except G59, G61, G62, and G64) of the evaluated accessions had little to no root constriction, 34.4% (G1, G3, G4, G5, G6, G10, G11, G15, G22, G23, G30, G33, G38, G41, G43, G44, G45, G46, G47, G57, G60, and G64) had white or cream external root color, 5% (G23, G36, and G40) had yellow root pulp color, 31.3% (G2, G9, G12, G13, G15, G24, G28, G29, G33, G34, G35, G36, G48, G49, G50, G52, G53, G55, G59, and G63) had white or cream root cortex, and 46.9% (G7, G9, G11, G13, G16, G20, G25, G26, G29, G32, G35, G37, G39, G40, G41, G42, G43, G45, G46, G48, G49, G50, G51, G53, G54, G55, G56, G57, G58, and G61) had ease of removing the root cortex.

Phenotypic characterization and evaluation could make the management and the use of several gene banks more efficient [34, 35]. Hence, the characterization and evaluation of plant genetic resources indicate the procedures used to assess, classify, distinguish, identify, avoid duplication, and efficiently utilize accessions. As a result, Francisco et al. [36] estimated the genetic variability in 16 sweet cassava accessions using 33 qualitative characters and found high divergence among the accessions. In Vietnam, cassava characterization and evaluation have been carried out for genetic variability and to classify seven cassava accessions using 20 morphological descriptors [37]. They found variability among accessions and classified them according to their morphological characteristics. Similarly, Zago et al. [12] evaluated 158 cassava accessions in Brazil to assess genetic variability using 38 qualitative characters, and the 37 characters showed variation among the tested accessions.

In the present study, several morphological characteristics revealed sufficient variation within the tested accessions. Among the characters, the shape of the central leaflet, leaf retention, petiole color, branching habit, the color of the stem epidermis, the color of the stem exterior, the external color of the storage root, and the color of root pulp were found to be the most variable factors in the evaluated cassava accessions towards frequency distribution. The variability of characters as indicated by frequency distribution in cassava agrees with the findings of Brice et al. [28], who found petiole color, branching habit, the color of root pulp, the color of the stem exterior, and plant shape to be the major discriminating characteristics. The other study was conducted by João Afonso et al. [38], who observed high variation in the foliar scar prominence, the cortex stem color, the petiole color, and the shape of the lobe. Carine et al. [39] reported that petiole color, branching habit, the shape of the plant, flowering, shape of the central leaflet, the orientation of the petiole, the color of end branches, and the color of the root cortex were observed to have high variability and offered possibilities for selection. Furthermore, the study reported by Nadjiam et al. [40] stated three variants of the stem exterior color, the stem epidermis color, and branching habit. However, these authors also found that the cortex stem color, the measurement between leaf scars, and the foliar scar prominence did not show variability among the evaluated 59 cassava accessions. This may be due to genetic similarities among accessions.

3.2. The Shannon–Weaver Diversity Index (H′)

The Shannon–Weaver diversity index (H′) was used to calculate the diversity of cassava accessions based on the frequency distribution of 30 qualitative characters and phenotypic classes [41]. In this aspect, the H′ value for most observed phenotypic characters exhibited a normal level of diversity among tested cassava accessions, which ranged from 0.24 for root constriction to 1.47 for petiole color, with a mean value of 0.84 (Table 3). As adopted by Islam et al. [42], H′ is classified as low (H′ < 0.50), medium (H′ = 0.50–0.75), and high (H′ ≥ 0.75). Based on this classification, four characters were categorized as low diversity, such as the color of the stem cortex, the presence or absence of fruit and seed, and root constrictions (Table 3). Additionally, eleven characters had medium genetic diversity, while the remaining fifteen characters had high genetic diversity.

A low level of diversity may indicate the narrow genetic base of the plant and be more frequent than others [43], while a high H′ value shows a relatively high level of diversity and an even distribution of the landraces [41, 44]. Moreover, the overall mean H′ value of 0.84 defined the presence of high phenotypic diversity among the tested cassava accessions. Thus, hybridization among accessions could produce the best hybrids and desirable segregants. This result is in agreement with the work of João Afonso et al. [38], who found a high H′ value for petiole color, stem exterior color, and color of the root bark.

3.3. Multiple Correspondent Analysis

The multiple correspondent analyses (MCA) of the qualitative characters are presented in Table 4. The first three dimensions of MCA were found to be responsible for up to 39.39% of the overall variation among the accessions (Table 4). The petiole color, leaf retention, the shape of the plant, pubescence on apical leaves, the external color of the storage root, and the color of root pulp were major contributors to the first dimension in multiple correspondent analysis (Dim1), which explained 20.77% of the total variation (Tables 4 and 5). The second dimension in multiple correspondent analysis (Dim 2) explained 9.98% of the total variation and was primarily related to the contrasting effects of root epidermis texture, lobe margins, root cortex color, leaf vein color, and petiole orientation (Tables 4 and 5). This finding was supported by those of Brice et al. [28], who characterized 89 accessions using 19 morphological characters, where the color of the apical leaves, the petiole color, flowering, and the yellow pulp color were the major contributing variables. In another study, Agre et al. [45] assessed the degree of genetic diversity of 116 elite cassava collections in Benin using 41 qualitative traits, while the apical leaf color, the shape of the plant, leaf retention, and constriction of the root are the leading contributors to the total variability.

3.4. Cluster Analysis

The clustering analysis results are presented in Figure 1, and the four divergent clusters’ formation was performed using the criteria with a pseudo-F and a pseudo-t-squared, which had values of 7.9 and 2.4, respectively; the maximum increase occurred at this point (Figure S1). The distribution of the 64 accessions (Table 6 and Figure 1) showed that 52 accessions were in cluster I, 6 in cluster II, 5 in cluster III, and 1 in cluster IV. The accessions grouped under cluster I have mainly been identified by the purplish green color of apical leaves, the absence of pubescence on apical leaves, outstanding leaf retention, greenish red petiole color, leaf lobe number of seven, smooth lobe margins, the horizontal orientation of the petiole, the light green stem cortex color, the white color of root pulp, and the pink color of root cortex (Tables S1, Table 6, and Figure 1). Additionally, the study found that three accessions, G23, G36, and G40, had roots with a yellow pulp color under this cluster (Table S1). It is interesting to note that one of the elements determining the root pulp’s value is its color. In cassava commercialization, it is preferable to use the pulp’s yellow color since it denotes a high content of carotenoids, which are precursors to vitamin A and are crucial for both human and animal diets [33, 46].

Under cluster II, six accessions were grouped together by having the difficult cortex removed or peeled, the cream color of root pulp, and the yellow color of root cortex, while cluster III was represented by the light brown external color of storage root, the white color of root pulp, the white or cream color of root cortex, and the easy cortex removed or peeled (Tables S1, Table 6, and Figure 1). Finally, one accession was classified as cluster IV due to the following characteristics: reddish green in less than half of the lobe, leaf vein color, red petiole color, inclined upwards orientation of the petiole, zigzag growth habit of the stem, the dark brown external color of the storage roots, the pink color of root pulp and cortex, difficulty in cortex peeling, and rough texture of the root epidermis (Tables S1, Table 6, and Figure 1).

Variations in morphological characters have been considered simple indicators of genetic variability in crop species and varieties. Cluster analysis is used to categorize accessions that are alike into one group and others into a different one [20]. Based on unweighted pair group methods, the 64 accessions were classified into four clusters, and the accessions represented in each cluster varied from 1 in cluster IV to 52 in cluster I, indicating that the studied cassava accession has a good chance of improving a character’s interest through effective selection. The distribution of accessions (Tables S1, Table 6, and Figure 1) showed that there were 52 (81.25%) accessions in cluster I (39 from the Hawassa, 3 from the Jimma, and 10 from the Nigeria); 6 (9.38%) accessions in cluster II (3 from the Hawassa, 1 from the Jimma, and 2 from Nigeria); 5 (7.81%) accessions in cluster III (2 from the Hawassa and 3 from the Nigeria); and 1 (1.56%) accessions in cluster IV from the Jimma. Accessions from the same or different sources are grouped into different clusters, implying that the genetic make-up of the accessions differs.

The clustering analysis result is consistent with a similar study in Côte d'Ivoire, where a total of 89 accessions of cassava were characterized using 19 qualitative characters and were classified into three groups [31]. In Benin, a total of 116 accessions were classified, by Agre et al. [45], into six clusters using 41 qualitative descriptions, and in Brazil, 45 cassava cultivars were classified, by Nadjiam et al. [40], into 5 clusters based on 36 qualitative characteristics. All those studies reported the presence of wide variability among tested accessions.

4. Conclusion

The aim of this study was to characterize and evaluate cassava accessions in order to provide useful information for breeding programs and conservation. In this regard, the evaluated accessions showed high variability for the shape of the central leaflet, branching habit, leaf retention, petiole color, color of the stem epidermis, the color of the stem exterior, external color of the storage root, and color of root pulp towards frequency distribution analysis. Thus, the various organ colors had a great role in identifying accessions. The Shannon–Weaver diversity index (H′) value for most observed phenotypic characters has exhibited an optimum level of diversity among accessions. The multivariate statistical analysis allows for identifying and grouping the accessions into different categories for various characters individually. As a result, the first two dimensions in the multiple correspondent analyses revealed 30.75% total variability, which was mainly related to 11 major characters. Thus, those characters are considered the most relevant for use in describing cassava accessions. In the cluster analysis, the 64 accessions are classified into four groups, with the number of accessions shared by each cluster being not uniform and varying from 1 in cluster VI to 52 in cluster I. This study could also facilitate breeders’ identification and classification of desired features for the cassava storage root from both a genetic improvement and an agronomic point of view. For instance, accessions G23, G36, and G40 are identified for optimal nutritional content. Finally, this study confirmed the existence of sufficient genetic variability for the characteristics evaluated, which could be attributed to the dissimilar genetic backgrounds of the evaluated accessions. Thus, the observed variation could be useful for a breeding program to develop cassava cultivars with desired root characteristics and conservation.

Data Availability

The datasets that support the findings of this study are available from the corresponding author upon request.

Conflicts of Interest

The authors declare that they have no conflicts of interest.

Authors’ Contributions

Berhanu Bilate carried out the field work, performed the statistical analysis, and drafted the manuscript. The rest of the authors coordinated the study, supervised fieldwork, and contributed to the writing of the manuscript. All authors have read and given their approval for the final manuscript to be submitted.

Acknowledgments

The universities of Jimma and Wolita Sodo are acknowledged by the authors for providing the funding for the study. The authors are also pleased to acknowledge Jimma and Hawassa Agricultural Research Centers and International Institute for Tropical Agriculture for providing planting materials used in the study. The research work was funded by Jimma and Wolita Sodo Universities.

Supplementary Materials

Table S1: The 64 cassava accessions and their score of qualitative characteristics. Figure S1: Criteria for the number of clusters based on a pseudo-F and pseudo-t-squared values. (Supplementary Materials)

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Copyright © 2023 Berhanu Bilate Daemo et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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