©Haramaya University, 2021
ISSN 1993-8195 (Online), ISSN 1992-0407(Print)

East African Journal of Sciences (2021) Volume 15 (1) 1-16

Licensed under a Creative Commons *Corresponding Author: wmanamno@gmail.com.
Attribution-NonCommercial 4.0 International License.

Genetic Variability and Correlation of Traits among Progenies of Potato Crosses in
Ethiopia

Manamno Workayehu 1*, Wassu Mohammed 2, and Tesfaye Abebe 3

1Adet Agricultural Research Center, P.O. Box 8, Bahir Dar, Ethiopia
2School of Plant Sciences, Haramaya University, P.O. Box 138, Dire Dawa, Ethiopia
3Holetta Agriculture Research Center, Holetta, Ethiopia

Abstract
Background: Investigating the causes and magnitude of genetic variation in segregating potato
population that derived from crossing is vital to know the genetic consequences of hybridization and
improve potato varieties. However, very little effort has been carried in creating local population
through crossing and genetic information on created potato population.
Objective: The study was conducted to assess genetic variability and correlation among traits in
locally created potato crosses population.
Materials and Methods: A total of 81 genotypes were evaluated for 18 traits in a 9 x 9 simple lattice
design. Data collection and analysis was done from sixteen plants or central rows.
Results: The results revealed highly significant variations for all traits except proportion of medium
tuber size and specific gravity of tubers indicating the existence of genetic variability among
population. Marketable and total tuber yields variability of tested genotypes ranged from 2.51 to 55.62
t ha -1 and 10.82 to 58.31 t ha -1, respectively. The phenotypic and genotypic coefficients of variation
ranged between 4.67 to 92.88% and 3.25 to 73.5%, respectively. Heritability in broad sense and
genetic advance as percent of mean also ranged from 28.81 to 91.64% and 4.65 to 90.33%,
respectively with less influence of environmental fluctuations. Total tuber yield had positive and
significant phenotypic and genotypic correlations with stem height, tuber yield per plant, tuber
number per plant, average tuber weight and marketable tuber yield. This indicated that the traits are
heritable with governing of additive gens for effective selection.
Conclusion: The range and mean values of the variables obtained suggest the existence of sufficient
variability among the tested and possibility of wide genetic base creation for improving potato
population using locally created genotypes. Hence, promising genotypes with desirable traits could be
recommended to produce new variety or use as parental lines for future breeding program.

Keywords: Broad sense heritability; Correlations; Genetic advance; Genotypes; Tuber yield

1. Introduction

Potato (Solanum tuberosum L.) is a highly heterozygous
tetraploid plant, originating in South America. It is a
tuber bearing crop belonging to the family
Solanaceous and has 48 chromosomes. Potato is the
world’s third most important food crop in overall
production after rice and wheat, and is a food security
crop in Ethiopia (Devaux et al. 2014). It is mainly
used as vegetable and available in the market
throughout the year with reasonable price and has
great importance in rural economy of the country as
compared to other vegetables crops in Ethiopia.
In Ethiopia, potatoes are mostly cultivated in the
central, north western, southern, and eastern parts of
the country (Semagn Asredie et al., 2016). The crop
has potential for improving the livelihoods of
millions of smallholder farmers in the high lands area
of the country. The potential for higher yield per unit
area, early maturity, and excellent food value give the
potato crop greater potential for improving food
security, increasing household income, and reducing
poverty than other crops (Semahagn Asredie et al.,

2015). In 2018/19, the area under potato production
of Maher season in the country was about 73,677.64
hectares with an average tuber yield of 14.18 t ha-1
(CSA, 2019). This is relatively low, especially when
considering the favorable climate at higher elevations,
soils, and irrigation potential in Ethiopia. The main
production constraints are related to narrow genetic
basis and susceptibility to diseases among varieties.
Ethiopia’s tremendous variation in altitude,
temperature, rain fall, soil type and ecological settings
also give rise to the need for a wide range of varieties,
which are not likely to be provided by existing
breeding programs (Semahagn Asredie et al., 2015).
So, continuously developing new cultivars through
crossing is needed for a sustainable increase in potato
variability and production under the present
environmental change and high human population
growth (Bradshaw, 2006).
Ethiopia has more than 36 potato varieties that are
approved for cultivation to address the production
problems in the country (MoA, 2016). These
improved varieties were developed from introduced
germplasms mainly from International Potato Center

https://link.springer.com/article/10.1007/s12230-016-9543-3#CR7

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

(CIP) either in the form of true potato seeds or
clones and varieties imported to the country through
technology shopping programs for adaptation trials
and subsequent registration as a variety
(Gebremedhin Woldegiorgis et al., 2013).
Creating genetic variability in tetraploid potato crop
through hybridization in the country is limited due to
too much dependence on CIP materials (Getachew
Assefa et al., 2016). Because of this constraint most
smallholder farmers are still growing old varieties that
are low yielding and disease susceptible. Improving
productivity of the crop through hybridization is
necessary to develop varieties which are adaptable to
a wide range of environments (Semahagn Asredie et
al., 2015). Ethiopian local potato varieties are more
heterozygous than the CIP genotypes that are
cultivated in Ethiopia (Semahagn Asredie et al., 2016).
Varieties developed from crossing of existing local
varieties are more likely to be adapted to local
growing conditions. Therefore, creating genetic
diversity by conventional breeding using locally
adapted genotypes (existing local land races and
released varieties) in the country is required to reduce
dependency on foreign materials and develop climate
smart potato varieties which are widely adaptable and
fulfill growers’ preferential traits.
Studying the causes and magnitude of variation
from segregating population or genotypes derived
from conventional crossing is vital in understanding
the genetic consequences of hybridization to develop
better potato varieties (Mehboob et al., 2016).
Evaluating crossing products in different generations
of selection can also help to estimate genetic
parameters such as heritability to identify the best
breeding strategy (Antonio et al., 2012).
Genetic variability in potato genotypes that were
introduced from CIP has been studied and variability
in segregation was reported by many researchers like
Addisu Fekadu et al. (2013); Abraham Lamboro
(2014); Getachew Assefa et al. (2016); Wassu
Mohammed (2015) and Tesfaye Abebe et al. (2012).
But not much effort has been made in a country to
study the genetic variability and improve varieties by
generating genotypes through crossing. Hence, the
present investigation was conducted to evaluate the
extent of variability, heritability, genetic advance, and
correlation of traits in populations of cross Jalene x

AterAbaba, Belete x Aterababa and Gera x Shenkola
varieties for different yield and its related traits.

2. Material and Methods
2.1. Description of the Study Site
The experiment was conducted at Adet Agricultural
Research Station during the main growing season in
2018. Adet Agricultural Research Center is located at
the longitude ranging from 37° 28’ 38’’ to 37° 29’ 50’’
E and latitude ranging from 11° 16’ 19’’ to 11° 17’
28’’ N in northwestern highlands of Ethiopia with an
average altitude of 2240 meters above sea level
(Andualem Wolie et al., 2013). The mean annual total
rainfall during the growing season was 1432 mm with
the average minimum and maximum temperatures of

10.81 and 25.55 ℃, respectively.

2.2. Description of the Experimental Materials
and Design
The experimental materials comprised 81 genotypes.
From this, seventy-five genotypes were offspring
produced from biparental crossing of Ethiopian
potato varieties by Adet Agricultural Research Center
in 2015. In addition, five high yielding parent
varieties, namely, Belete, Aterababa, Gera, Shenkola
and Jalene and Dagim (the latest registered variety)
was included as a standard check for comparison in
this experiment (Table 2).
The experiment was laid out as a 9 x 9 simple lattice
design. Each genotype was planted in a gross and net

plot size of 1.5 m x 3m which contained two rows in
plot and twenty plants per plot. Medium-sized and
well sprouted potato tubers were planted at the
spacing of 75 cm between rows and 30 cm between
plants. The recommended dose of fertilizer was
applied at a rate of 81/69 N/P

2
O

5
per hectare. The

whole phosphors fertilizer was applied during
planting, but N source applied in split at planting, 2

weeks after emergence and at flowering at equal
1

rate. Earthing up was executed two times throughout
the entire growing period, one at 30 days and another
one at 60 days after planting. Fungicide chemical
(Ridomil) was applied once when symptom occurred
on experiment to control potato late blight disease.

Manamno et al. Genetic Variability among Potato Progenies

Table 1. Description of parental varieties for different traits.

S/N Parent Released Characteristics

1 Belete
CIP-393371.58

2009 Strong stems, high yield and dry matter, excellent stew quality, late
blight tolerance, early bulking, early maturing and good for French fries,
large tuber, white flower, and tuber skin color, very susceptible for
bacterial wilt, hollow heart, sometimes its shape become amorphous,
bitter taste than others

2 Jalene
CIP-37792.5

2002 Good dry matter and boiling quality, early maturing, early flowering,
white tuber skin and flower color, late blight susceptible shallow eye
depth, good taste

3 Gera
KP-90134.2

2003 Long storage life, strong stems, high yield, late blight tolerant and well
adapted to dry areas, very high deep tuber eye, white flower and tuber
skin, round tuber shape

4 Shenkola
KP-90134.5

2005 Good taste and boiling quality, white flower and tuber skin color, good
yield, oval tuber shape, shallow eye depth, good taste, has tall stem

5 Aterababa Local Good canopy cover, stew quality, taste, early maturing, round shape,
high dry matter, good market demand and storage quality, late blight
susceptible, bacterial wilt resistant than others, pink tuber skin and
flower color, round tuber shape

Note: Descriptions are organized from Tesfaye Abebe et al. (2013) and Semahagn Asredie et al. (2015).

Table 2. List of potato offspring and parents with standard check variety used in the experiment.

Trt Genotype Trt Genotype Trt Genotype Trt Genotype

1 J x A.277 22 J x A.42 43 J x A.27 64 B x A.248
2 B x A.153 23 B x A.15 44 Ge xSh.186 65 J x A.18
3 J xA.296 24 J x A.49 45 J x A.130 66 J x A.123
4 B x A.174 25 B x A.60 46 B x A.163 67 B x A.207
5 J x A.94 26 J x A.77 47 J x A.67 68 J x A.186
6 B x A.225 27 Gera 48 Shenkola 69 B x A.129
7 Ge x Sh.65 28 J x A.31 49 Ge x Sh.206 70 J x A.122
8 Belete 29 Ge x Sh.101 50 J x A.146 71 J x A.243
9 J x A.140 30 J x A.333 51 B x A.8 72 Ge x Sh.90
10 B x A.74 31 B x A.228 52 J x A.102 73 Ge x Sh.317
11 J x A.170 32 J xA.266 53 B x A.213 74 J x A.196
12 B x A.112 33 J x A.143 54 J x A.245 75 J x A.250
13 J x A.21 34 J x A.326 55 J x A.345 76 J x A.119
14 B x A.184 35 Dagim 56 B x A.201 77 J x A.246
15 B x A.164 36 J x A.188 57 Ater ababa 78 J x A.165
16 J x A.120 37 J xA.60 58 J x A.135 79 Jalene
17 J x A.187 38 B x J.16 59 B x A.603 80 B x A.97
18 B x A.44 39 J x A.34 60 J x A.201 81 Ge x Sh.96
19 J x A.39 40 Ge x Sh.319 61 B x A.55
20 B x A.198 41 B x A.140 62 J x A.9
21 Ge x Sh.29 42 J x A.23 63 Ge x Sh.100

Note: Trt = Treatment, J x A = Jalene cross Ater abab, B x A = Belete cross Ater ababa, Ge x Sh = Gera cross Shenkola, Dagim
= Standard check variety and numbers followed crosses indicated the code of genotype (experimental material).

2.3. Data Collection and Analysis

2.3.1. Data collection

Observations were recorded on different traits such
as days to emergency (DE), days to flowering (DF),
days to maturity (DMA), main stem number (SN),
plant height (PH), tuber number per plant (TNP),
tuber yield per plant (TYP), very small tuber numbers
(VSN), medium sized tubers (MDN), large sized
tubers (LTN), tuber dry matter content (DM), tuber
starch content (SC), tuber specific gravity, average

tuber weight (AW), marketable tuber number (MTN),
marketable yield (MY), unmarketable tuber yield
(UNMY) and total tuber yield (TY). Some of
morphological qualitative traits like, flower color,
plant growth habit type, predominant tuber skin color
and flesh color, tuber skin type, eye depth and
number per tuber were recorded in all the entries as
per the standards and codes specified in potato
descriptors of AICRP on potato, Huaman et al.
(1977) during the peak of crop growth.

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

2.3.2. Data Analysis
The quantitative data were subjected to analysis of
variance (ANOVA), but descriptive statistics was
used to describe qualitative data by taking samples
from five plants for each trait. Means for significant
treatments were compared using Fisher ‘s protected
least significant differences (LSD) at 5% (P<0.05)
level of significance. The traits that exhibited
significant mean squares in ANOVA were further
subjected to genetic analyses. The phenotypic (σ2p),
genotypic (σ2g) variances, environmental variance (σ2e)
and the corresponding phenotypic (PCV) and
genotypic (GCV) coefficients of variation for each
trait were estimated following the method described
by Burton and Devane (1953). Genotypic variance

(σ2g) =
𝐺𝑀𝑆−𝑀𝑆𝐸

𝑟
, Phenotypic variance (σ2p) = σ2g +

σ2e. where, MSE =Mean square of error and
GMS=genotypic mean square. Broad sense
heritability estimates and genetic advance were also
calculated using the formula of Burton (1952) and
Johnson et al. (1955), respectively: h2b (Broad sense

heritability) (%) =
σ²g

σ²p
x100, where σ2g= genotypic

variance and σ2p=Phenotypic variance. Genetic
advance (GA) for each trait was computed using the
formula adopted by Johnson et al. (1955) as: GA=
GA=k.σp.h2b.Where, k= selection differential (k=2.06
at 5% selection intensity), σp= Phenotypic standard
deviation of the trait, h2b= broad sense heritability of
the character.

3. Results and Discussion
3.1. Analysis of Variance and Mean Performance
of Genotypes
Results of the analysis of variance (ANOVA) of 18
quantitative traits for the 81 genotypes showed
significant (P ≤ 0.01) differences among the tested
potato populations for all traits except proportion of
medium tuber size (%) and specific gravity (g/cm 3)
of tubers (Table 3). The presence of significant
differences among the genotypes obtained from three
biparental crosses and their parents suggested the
chance of obtaining better performing
offsprings/genotypes than their parents for the
different important yield and its related traits. The
genotypes are expected to be highly heterozygous in
which additive and non-additive gene actions and in
most case, both operate (Ross, 1986), (Arndt and
Peloquin, 1990). Therefore, the observed variations
among the potato genotypes could be exploited
through vegetative propagation.
The present findings agree with findings of Hajam
et al. (2018) who reported significant variations
among 38 genotypes for days to flowering, plant
height, tuber number per plant, tuber yield per plant
and average tuber weight. Hirut Betaw et al. (2017)

reported highly significant differences among 60
progenies of 32 families, 12 twelve parents and check
varieties for growth, physiological and tuber yield
related traits. Melito et al. (2017) also reported highly
significant difference for total tuber yield from
genotypes obtained biparental crosses. Zakerhamid
(2014) showed significant differences among 166
hybrids and two parents for plant height, main stem
number per plant, tuber weight per plant, average
tuber weight and tuber yield. Nickmanesh and
Hassanpanah (2014) evaluated 127 hybrids with their
two parents and reported significant differences for
days to flowering, plant height, tuber weight per
plant, and total tuber yield.
The genotypes/offsprings, parents and the check
variety observed 16 to 24, 42 to 60 and 88 to 102
days of 50% emergence, 50% flowering and 90%
maturity, respectively (Table 4). The 41 potato
offsprings of Jalene x Ater Ababa had lowest mean
days to 50% emergency. The 24 potato progenies of
Belete x Ater Ababa also showed lowest mean days to
50% flowering. While the 10 potato offsprings of
Gera x Shenkola manifested lowest mean days to
maturity. However, the individual genotypes, J x
A.277, J x A.296, J x A.187, J x A.135, J x A.201, J x
A.122, J x A.196, J x A.246 and J x A.165 showed
early emergency of plants than their parents, standard
check variety and other genotypes. Similarly, J x A.23
and J xA.130 genotypes observed early maturity (88
days) than other progenies, parents, and check
variety. A total of nine potato progenies showed early
maturity than the standard check variety in this study.
The main stem number and stem height ranged
between 2 and 9; and 32.27 and 73.33 cm,
respectively (Table 4). Potato offspring of Jalene and
Ater Ababa exhibited the highest mean of main stem
numbers whereas the maximum mean stem height
recorded for progenies of Gera and Shenkola.
Genotype B x A.164 showed maximum stem height
(73.33 cm) followed by Ge x Sh.319 (71.64 cm) but
more stem numbers recorded for genotype J x A.9.
These genotypes which had more main stems and tall
stem height can produce high tuber yield.
Other authors also found the performance of
crosses and parental varieties for phenological and
growth traits. Hajianfar et al. (2017) recorded main
stem numbers ranging between 3.33 and 4.29 and
stem height ranging between 52.33 and 64.77cm for
potato hybrids. Parmar et al. (2015) found plant
height ranging from 37.16 to 54.80 cm with a mean
of 42.29 cm and main stem number per hill ranging
from 1.93 to 3.93 with a mean 2.91 in potato
genotype. Biswas (2010) showed the range of plant
height from 15.0 to 57.0 cm with a mean 37.9 cm for
and 1 to 6 with a mean 2.1 for main stem numbers
for potato progenies.

Manamno et al. Genetic Variability among Potato Progenies

Table 3. Mean squares from the simple lattice ANOVA for traits of 81 potato genotypes.

Trait Rep
df =1

Blocks (adj.)
df =16

Genotype (unadj.)
df = 80

Genotype (adj.)
df = 80

Error (RCBD)
df = 80

Intra block (Error)
df = 64

RE of SL
Over RCBD

CV (%)

DE 32 3.49 8.14** 7.74** 2.5 2.25 103.61 8.36

DF 9.88 4.2 77.32** 73.56** 3.54 3.37 100.92 3.97

DMA 52.25 6.50 28.89** 28.76** 9.36 10.07 92.91 3.24

SN 2.47 0.82 3.12** 2.96** 0.93 0.96 96.99 25.78
SH 99.25 37.69 168.85** 150.79** 25.51 22.47 105.06 10.03
TNP 0.5 3.74 30.17** 25.12** 4.58 4.78 95.65 16.3

TYP 0.0005 0.007 0.09** 0.08** 0.008 0.008 96.79 14.21

AW 30.33 53.52 587.04** 565.85** 61.58 63.59 96.83 16.7

MTN 54267 1562.95 3916.31** 3360.49** 1185.46 1091.09 102.46 25.04

MY 210.12 12.72 200.13** 192.04** 16.4 17.33 94.68 16.55

UNMY 286.56 3.75 20.86** 18.77** 3.83 3.86 99.42 31.3

TY 5.89 10.92 189.95** 177.46** 14.97 15.99 93.66 16.69

VSN 17515 139.8 275.23** 265.25* 142.08 142.66 99.6 32.63

MDN 19781 190.98 210.36ns 191.5ns 155.2 146.26 101.37 24.43

LTN 50.72 48.9 335.45** 327.70** 52.12 52.92 98.48 31.36

DM 37.07 3.88 20.70** 19.61** 5.4 5.78 93.43 11.05

SG 0.14 0.0009 0.001ns 0.001ns 0.002 0.002 89.13 3.53

SC 34.86 3.25 16.43** 15.52** 4.68 5.04 92.89 14.48

Note: *and **Significant at P< 0.05 and P<0.01 probability levels, respectively. DE = days to emergency, DF = days to 50% flowering, df=degree of freedom, DMA = days to 90% maturity, SN = main
stem number, SH (cm) = stem height, TNP = tuber number per plant, TYP (g) = tuber yield per plant, AW (g) = average tuber weight, MTN = marke table tuber number, MY = marketable yield t ha-1,
UNMY = unmarketable yield t ha-1, TY = total yield t ha-1,VSN = very small tuber size in percentage, MDN = medium tuber size percentage, LTN = large size tuber percentage, DM = tuber dry matter
content(%), SG = specific gravity of tuber, SC = tuber starch content (%), CV (%) = coefficient of variation.

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

Tested genotypes showed 6 to 25 tuber number per
plant, 0.23 to 1.025kg tuber yield per plant,16.61 to
106.83g average tuber weight, 11 to 310 marketable
tuber number per net plot, 2.51 to 55.62 marketable
tuber yield, 0.33 to 24.9 unmarketable and 10.82 to
58.31ha-1 total tuber yield (Table 4). The result of
range and mean of marketable and total tuber yield
were greater than the findings reported by Getachew
Assefa et al. (2016), Wassu Mohammed (2014), Wassu
Mohammed and Simret Burga (2015), Addisu Fekadu
et al. (2013) and Tesfaye Abebe et al. (2012) from
tested Ethiopian cultivated varieties and non-
registered CIP genotypes. The greater number of
tubers per plant and marketable tuber number per
plot were 25 and 246.5 for genotypes J x A.170 and J
x A.18. On the other hand, the highest and
significantly different tuber yield per plant,
marketable tuber yield and total tuber yield noted
from genotype B x A.164, followed by J x A.119. In
contrast, the highest significant unmarketable tuber
yield 24.9 t ha-1 was recorded for genotype Ge x
Sh.29, followed by Ge x Sh.206 13.59 t ha-1 due to
very deep eye depth and cracking of tubers. This may
show dominance of maternal inheritance traits which
Gera variety (female parent) imparts very deep eye on
tubers.
Generally, in the current study, about 40% potato
offspring exhibited significantly higher total tuber
yields than the standard check Ethiopian variety.
Single genotype B x A.164 and J x A.119 showed
total tuber yield advantage of 209.17% and 133.96%,
respectively, over the standard check and about
36.7% and 3.5% yield advantage over Belete (high
yielding registered variety) so far in Ethiopia. In a
similar kind of study, Melito et al. (2017) found that
48% of the genotypes from a total of 27 populations
which produced from seven biparental
crosses/families that showed higher total tuber yield
compared to the standard check.
The highest mean tuber number per plant and
marketable tuber number recorded from biparental
crosses of Jalene and Ater ababa whereas the
maximum mean of tuber yield per plant, average
tuber weight and unmarketable tuber yield found in
crosses of Gera x Shenkola variety (Table 4). In
addition, potato genotypes (offsprings) produced
from Belete cross with Ater Ababa manifested
highest mean for marketable tuber yield and total
tuber yield. This may be due to over dominance of
maternal effect which Belete (female parent) is high
yielder variety in the country.
Tuber size distribution such as proportion of small
tubers, medium tubers and large tubers ranged from
5.86 to 59.07; 26.8 to 81.08; and 0.6 to 55.67%,
respectively (Table 4). The maximum mean
percentage of small tubers and large tubers were
noted from progenies of Jalene and Ater Ababa (41)
and Gera and Shenkola accordingly. The maximum
proportion of small tubers (59.07%) showed by
genotype J x A.333 whereas, genotype Ge x Sh.206
and parental varieties such as shenkola and belete

variety exhibited significantly highest percentages of
large tubers.
Proportion of quality traits (tuber dry matter and
starch content) ranged from 13.78 to 29 and 8.28-
21.84% (Table 4). These results agree with the
findings of Garnica et al. (2012) who reported tuber
dry matter and starch content (%) that ranged from
22.1 to 28.06 and 15.86 to 21.51%, respectively. But
higher than explained by Tesfaye Abebe et al. (2012)
who evaluated 25 released potato varieties and the
varieties overall values ranged from 17.65 to 26.70%
dry matter content and 9.75 to 17.82% for total
starch content under Adet environment. Potato
offsprings of Belete and Ater Ababa produced higher
mean tuber dry matter yield and tuber starch content
(%) than other biparental crosses. Significantly lower
tuber dry matter and starch content (%) were
recorded from crosses of Gera and Shenkola variety
by genotype Ge x Sh.186. Individual genotypes B x
A.97 and J x A.266 showed higher tuber dry matter
yield and starch content than all other clones. The
results may be due to positive correlation of dry
matter content with total starch content (r=1**) as
observed in this study and suggests that continuous
evaluation of these crosses may result in a higher
chance of obtaining genotypes with high tuber quality
traits. A total of forty (40) potato offsprings and four
parental varieties produced the maximum percentages
of tuber dry matter yield and starch content than the
standard check variety.
The present findings are also supported by results
of previous workers. Addisu Fekadu et al. (2013)
reported availability in tuber size distribution that
ranged from 4.6 to 56.67% for small sized tubers,
27.80 to 49.00% for medium sized tubers and 0.5 to
65.7% for large sized tubers. Melito et al. (2017) also
reported the possibility of obtaining many genotypes
with better tuber yield from bi-parental crosses.
Similarly, Hajianfar et al. (2017) reported as high as
11.96, 41.22 and 37.19 t ha-1 tuber number per plant,
total tuber yield, and marketable yield, respectively,
and tuber dry matter up to 26.7% in hybrids of
potato genotypes. In addition, Luthra et al. (2017)
observed the range from 6.23 to 10.34 for tuber
number per plant, 20.2 to 36.6 for marketable yield,
23.7 to 38.8 for total yield t ha-1 and 13.9 to 18.5%
for dry matter from potato progenies. Additionally,
Parmar et a l. (2015) showed tuber yield per plant
249.15 to 511.07gm and average tuber weight 48.53
to 98.2gm among 32 potato progenies. Furthermore,
Nizamuddin et al. (2010); Zakerhamidi (2014) and
Nickmanesh and Hassanpanah (2014) reported a
wide range of tuber yield in progenies of two varietal
crosses. Feltran et al. (2004) reported high dry matter
and starch content ranged from 15.7 to 22.4% and
56.7 to 71.4%, respectively, in 18 potato cultivars.
In family base, the maximum proportion in 41
potato offspring obtained from crossing Jalene with
AterAbaba showed semi-erect growth habit (39.02%),
light pink flower color (80.49%), brown tuber skin
color (29.27%), yellow tuber flesh color (44.34%),
medium tuber eye depth (39.02%), smooth skin type

Manamno et al. Genetic Variability among Potato Progenies

(58.54%), oval tuber shape (39.02%) and intermediate
(5<20) eye numbers(39%). On other hand, most
proportion of potato offsprings/genotypes derived
from Belete and Ater Ababa contained decumbent
growth habit (45.83%), light pink flower (58.33%),
brown tuber skin color (37.5%), yellow tuber flesh
color (41.67%, medium tuber eye depth (54.16%),
smooth tuber skin type (70.8%), round and long
tuber shape (60%) and inter mediate tuber eye
number (91.67%). However, the proportion of the
genotypes generated from Gera cross Shenkola
observed decumbent (45.83%) growth habit, light,
and intense purple flower color (100%), light brown
(30%) and white tuber skin color (50%), light yellow
tuber flesh color (40%), light yellow tuber flesh color
(40%) for each) and oval tuber shape (70%) in this
study.
According to Kumar et al. (2018) white-flesh-
colored potatoes are low in carotenoids
(<100μg/100 g fresh weight) whereas the carotenoids
content of yellow-fleshed varieties is higher (about
560μg/100 g FW). Intense yellow to near orange
flesh color is associated with carotenoid
concentrations >2000μg/100g. Based on this
information, about 33 from a total of 75 potato
progenies showed white and cream flesh color which
is low in carotenoids, while other genotypes
contained yellow and different flesh color which had
high carotenoid content. Cultivars which possess
either shallow or medium eye depths are perfect to
reduce losses during peeling and trimming.
Therefore, the results of crossing Belete and Ater
Ababa produced large proportion of skin type highly
desired for any French frying or processing industries
and as such considering these parental materials
seems essential to producing clonal population
targeting processing industries. Hence, genotypes in
this study showed difference by morphological
qualitative traits not only among all families but also
with in biparental crosses.

3.2. Genetic Variability Components
3.2.1. Phenotypic and genotypic coefficients of
variations
The variability components (phenotypic and
genotypic variances and coefficient of variations,
heritability in broad sense and genetic advances as
percent of mean) were estimated for agro
morphological traits (Table 5). The estimates of
genotypic and phenotypic coefficient of variations
were computed in the range between 3.25 to 51.41
and 4.67 to 60.28%, respectively. The lowest and the
highest values were computed for days to maturity
and number of large tubers, respectively. According
to Siva Subramanian and Menon (1973), PCV and
GCV values roughly more than 20% are high,
whereas values less than 10% are low and values
between 10 and 20% to be medium.
Correspondingly, days to flowering, plant stem
height, average tuber weight, marketable tuber
number, marketable tuber yield and total tuber yield
exhibited high phenotype and genotypic variance

indicating greater scope of selection for the
improvement of these characters. Similarly, Haydar
(2009) reported the maximum genotypic and
phenotypic variance for plant height. Hajam et al.
(2018) also observed highest GV and PV for average
tuber weight, tuber yield per plant and total tuber
yield, and in addition, Benavente and Pinto (2012)
found high genotypic variance for total tuber yield
from among families and within families in 30 potato
genotypes which produced by biparental crossing.
Maximum phenotypic coefficients of variation
(PCV) was computed for main stem number, tuber
number per plant, average tuber weight, tuber yield
per plant, marketable tuber number, marketable yield,
total tuber yield, tuber starch content and percentages
of small tubers and large tubers whereas highest
genotypic coefficients of variation (GCV) had
observed for traits such as main stem number, tuber
number per plant, average tuber weight, tuber yield
per plant, marketable tuber number, marketable yield,
total tuber yield, percentages of small tubers and large
tubers. In line with this study, Ozturk and Yildirim
(2014) reported high genetic coefficient of variation
(GCV) for total yield (26.2 %), plant height (21.2 %)
and average tuber weight (20.4 %) for genotypes. The
characters having high GCV indicate high potential
for effective selection (Burton, 1957). Characters with
high genetic variability and genetic advance are also
important for selecting the desirable parents (Biswas
et al., 2005).
High estimates of phenotypic (PCV) and genotypic
coefficients of variation (GCV) were observed for
main stem number, tuber number per plant, average
tuber weight, tuber yield per plant, marketable tuber
number, marketable tuber yield, total tuber yield,
small tuber, and large tuber percentage. The result
was agreed with authors such as Hajam et al. (2018),
noted highest GCV and PCV for traits number of
main stems, total tuber yield and tuber yield per plant.
The traits which exhibited high estimates of
genotypic and phenotypic coefficient of variations
had high probability of improvement through
selection to develop new variety (Singh, 1990).
Moderately high phenotypic (PCV) and genotypic
coefficients of variation (GCV) were computed for
plant stem height, days to 50% flowering and tuber
dry matter. Moderate phenotypic coefficients of
variation (PCV) were also noted for days to 50%
emergency. Abraham Lamboro (2014) reported
moderately high PCV and GCV for plant stem
height, percentage of small tubers and days to
emergence. Low estimates of PCV and GCV were
obtained for days to maturity. This agrees with
Kameshwari (2015) who reported the lowest GCV
and PCV for days to maturity. The lowest GCV and
PCV suggested that selection for the desired
character based on phenotypic expression of
genotypes might not be effective in attaining the
desired genotypes due to the highest masking of
factors to express these traits. Singh (1990) also
reported low GCV and PCV values for dry matter
content.

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

Table 4. Average family mean values (range) of evaluated genotypes for quantitative traits.

Family No. of clone Range DF MA SN SH TNP TYP ATW MTN MY UNM TY VSN LTN DM SC

J x A 41 Min 44 88 3 33.1 7 0.23 16.61 66 6.58 0.33 10.8 6.25 0.6 14.13 8.59
Max 53 101 9 66 25 0.97 71.38 247 43.8 8.19 44.1 59.1 40.29 27.68 20.67
Mean 48 95 5 49.2 14 0.59 42.27 144 23.3 3.7 27.0 30 18.14 21.18 14.88
B x A 24 Min 42 88 2 32.3 6 0.27 26.45 75 8.73 1.05 11.7 5.86 3.2 15.63 9.93
Max 55 102 6 73.3 23 1.03 78.1 226 55.6 7.65 58.3 42.5 49.78 29.00 21.84
Mean 46 95 4 46.6 12 0.62 53.55 130 26.9 3.4 30.3 23.3 27.23 21.73 15.36
Ge x Sh 10 Min 46 90 3 41.3 8 0.3 35.11 13.5 2.51 0.58 16.8 13.2 12.6 13.78 8.28
Max 60 99 7 71.6 17 1.03 103.3 193 33.9 24.9 45.7 52.2 55.47 25.48 18.7
Mean 50 94 5 53.9 12 0.63 56.49 118 21.7 7.51 29.2 27.2 27.61 21.57 15.22
Jalene 1 Mean 54 93 4 50.2 15 0.6 40.96 135 23.0 4.41 27.4 36.3 17.81 18.15 12.18
Aterababa 1 Mean 45 91 4 48.1 13 0.49 39.32 108 17.1 5.34 22.4 34.9 13.19 22.18 15.76
Belete 1 Mean 48 99 2 52.9 8 0.88 106.8 111 41.0 1.66 42.6 13.2 55.67 24.3 17.66
Gera 1 Mean 52 99 2 65.8 13 0.89 72.87 157 37.1 2.23 39.3 14.3 41.31 20.98 14.69
Shenkola 1 Mean 55 101 2 66.6 10 0.82 82.11 127 36.0 1.6 37.6 16.9 55.59 23.75 17.17
Dagim 1 Mean 48 92 2 57.4 10 0.43 48.94 105 16.4 2.47 18.9 20.4 31.86 21.3 14.99
G.mean 48 95 4 49.3 13 0.6 48.74 135 24.5 3.9 28.0 27.0 23.0 21.47 15.14

Note: J x A = Jalene cross with Ater ababa, B x A = Belete cross with Ater ababa, Ge x Sh= Gera cross with Shenkola, Dagim = standard check variety, Min=minimum, Max = maximum,
G.mean=Grand mean, DE = days to emergency, DF = days to 50% flowering, DMA = days to 90% maturity, SN = main stem number, SH (cm) = stem height, TNP = tuber number per plant, TYP (g)
= tuber yield per plant, AW (g) = average tuber weight, MTN = marketable tuber number, MY = marketable yield(t ha -1), UNMY = unmarketable yield (t ha-1), TY = total yield(t ha-1),VSN = very
small tuber size percentage, MDN = medium tuber size percentage, LTN = large size tuber percentage, DM=tuber dry matter content (%), SG = specific gravity of tuber, and SC = tuber starch content (%).

Manamno et al. Genetic Variability among Potato Progenies

On this study, the PCV values were slightly higher
than their corresponding GCV values for all the
characters considered which reflect a little influence
of environment on the expression of characters. This
agrees with Getachew Assefa et al. (2016) and Biswas
(2005) reported high phenotypic variances than
genotypic variances for growth and yield traits in
genotypes.

3.2.2. Heritability and genetic advance
Estimate of heritability in broad sense ranged from
28.81 to 91.64% for percentages of small sized tubers
and days to flowering, respectively (Table 5). As
suggested by Pramoda (2002), h2b estimates is
categorized as low < 40%, medium 40-59%,
moderately high (60-79) and very high (>80%). Based
on this category, very high heritability estimate was
noted for days to flowering (91.64%), tuber yield per
plant (80%) and average tuber weight (80.45%)
indicating the selection of genotypes for such
characters could be easy and may lead to the
improvement of the mean values in selected
genotypes for those traits. Presence of high
heritability indicated that these traits are less
influenced by environmental fluctuations and
governed by the additive gene effects that are
substantially contributing towards the expression of
traits. Similarly, Ozturk and Yildirim (2014) reported
the highest board sense heritability’s for average tuber
weight, Moussa (2013) noted high estimates of
heritability for tuber yield per plant from 17 potato
genotypes comprising seven parents and their ten
crosses, and Haydar (2009) also reported high
heritability for tuber yield per plant and tuber
numbers per plant respectively from seven potato
parent and hybrids.
High heritability coupled with high genetic
advances was computed for average tuber weight
indicating the influence of additive gene effect on the
trait. Hence, these traits can be improved through
simple selection. Effectiveness of selection is
considered more reliable when heritability is coupled
with genetic advance. The result was in close

agreement with the findings of Moussa (2013) and
Biswas (2005) who reported that high heritability
value along with high genetic advance for average
tuber weight in potato hybrids.
Moderately high heritability was also ranged
between 72.75 and 77.86% for stem height, tuber
number per plant, marketable tuber yield, total tuber
yield and proportions of large sized tubers while
medium heritability showed for days to emergency,
days to maturity, marketable tuber number, tuber dry
matter and tuber starch content percentage. In line
with this study, Mishra et al. (2017) obtained
moderately high heritability for marketable yield
(77.1%), total tuber yield (76.8%) and tuber dry
matter from hybrids. Low heritability value noted for
small tuber size (28.81%). This indicates that
selection may be considerably difficult or virtually
impractical due to the masking effect of the
environment. High heritability accompanied with low
genetic advances was recorded for tuber yield per
plant. This suggested that the character was
influenced due to favorable influence of environment
or predominant effects of non-additive gene rather
than genotypes. Panse (1957) reported that low
heritability accompanied with genetic advance is due
to non-additive gene effects for the particular trait. As
stated, Panes and Sukhatme (1964), high heritability
values associated with equally high genetic advance is
chiefly due to dominance and epistasis.
The genetic advance as percent mean was
categorized as low (<10%), moderate, (10-20%) and
high (>20%) by Johnson et al. (1955). Based on these
categories, the highest genetic advance for tested
genotypes expressed as percentage of the mean
(GAM) showed for all traits ranging from 25.08 to
90.33% except for days to emergency, days to
maturity, percentages of tuber dry matter and tuber
starch content (Table 5). Traits which showed high
values of genetic advance might be due to additive
gene action. Similarly, Kameshwari (2015) reported
high genetic advance in percentage of mean for
average tuber weight and tuber yield per plant.

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

Table 5. Variability components for 16 traits of evaluated 81 genotypes and parental varieties.

Trait Range Mean σ2g σ2p GCV (%) PCV (%) Hb (%) GA (5%) GAM (5%)

Days to emergence 16-24 18.12 2.95 5.20 9.47 12.58 56.65 2.66 14.68

Days to flowering 42-60 61.00 36.97 40.35 12.72 13.28 91.64 11.99 25.08

Days to maturity 88-102 94.51 9.41 19.48 3.25 4.67 48.3 4.39 4.65

Stem height(cm) 32.27-73.33 49.34 73.19 95.66 17.34 19.82 76.51 15.42 31.24

Main stem number 2.0-9.0 3.80 1.08 2.04 27.23 37.43 52.9 1.56 40.79

Tuber number/plant 6.0-25.0 12.93 12.69 17.48 27.55 32.33 72.63 6.26 48.37

Average tuber weight(g) 16.61-106.83 48.74 261.72 325.32 33.20 37.01 80.45 29.89 61.33

Tuber yield /plant(kg) 0.23-1.03 0.60 0.04 0.05 32.63 35.79 80.00 0.37 61.26

Marketable tuber number 11-310 134.5 1412.61 2503.7 27.94 37.19 56.42 58.16 43.23

Marketable yield t ha-1 2.51-55.62 24.46 87.61 112.52 38.26 43.36 77.86 17.01 69.55

Total yield t ha-1 10.82-58.31 28.32 81.94 108.01 31.97 36.7 75.87 16.24 57.36

Tuber dry matter 13.78-29 21.47 6.23 14.47 11.62 17.72 43.02 3.37 15.7

Tuber Starch content 8.28-21.84 15.14 4.94 11.49 14.69 22.39 43.04 3.01 19.85

Small tuber 5.86-59.07 26.89 61.36 212.99 29.13 54.27 28.81 8.66 32.2

Large tuber 0.6-55.67 49.99 141.26 194.18 51.41 60.28 72.75 20.88 90.33

Note: σ2g = genotypic variance, σ2p = phenotypic variance, PCV (%) = phenotypic coefficient of variation (%) = genotypic coefficient of variation, Hb (%) = heritability in broad sense, GA = genetic advance,
and GAM = genetic advance as percent of mean.

Manamno et al. Genetic Variability among Potato Progenies

3.2.3. Phenotypic and genotypic correlations
The estimates of genotypic and phenotypic
correlation coefficients between total tuber yield and
all possible pairs of yield components are presented
in Table 6. Phenotypic and genotypic association of
days to emergency with days to flowering was
positive. This indicates that selection for this trait
may lead to early mature genotypes. Days to
emergency was negatively correlated with main stem
number, tuber number per plant and marketable
tuber number at phenotypic and genotypic levels but,
positively correlated with proportion of large tuber
size at both levels. Plant stem height positively
correlated with tuber yield per plant, tuber number
plant, average tuber weight, marketable tuber
number, marketable tuber yield, total tuber yield,
tuber dry matter, tuber starch content and proportion
of large tuber sized tubers at both correlations level
but, it correlated negatively with proportion of small
tuber size at genotypic level.
Total tuber yield showed positive genotypic and
phenotypic association with days to maturity, tuber
number per plant, average tuber weight, marketable
tuber number, tuber dry matter, tuber starch content
and proportion large tuber size but showed negative
association with proportion of small tuber sizes. Both
marketable tuber yield and total tuber yield were
positively correlated with plant stem height, tuber
yield per plant, tuber number per plant, marketable
tuber number and average tuber weight, tuber dry
matter, tuber starch content and proportion of large
tuber numbers at phenotypic level. These positive
correlations indicating that selection for improving
one character will lead to increase the other one
which is positively correlated with that character. In
contrast, marketable tuber yield and total tuber yield
were negatively correlated with proportion of small
tuber sizes. In similarly with Abraham Lamboro et al.
(2014) and Addisu Fekadu et al. (2013) who reported

highly significant negative correlation of total tuber
yield with proportion of small sized tubers at
genotypic and phenotypic level. Negative correlation
between two traits implies selection for improving
one character will likely cause decrease in the other
traits.
In present study the tuber dry matter had exhibited
strong phenotypic and genotypic correlation (r = 1)
with tuber starch content. This suggested that the
simple selection to improve one trait simultaneously
increase the second character. Similarly, highly
significant correlation was reported for dry matter
and tuber starch content by Wassu Mohmmad (2016)
who observed the correlation was near to perfect (r =
0.97 to 0.99) and Khayatnezhad et al. (2011) that
reported stronger positive and significant correlations
between starch content and dry matter content.
Generally, in most of traits genotypic correlation was
higher than phenotypic correlation indicating an
inherent association between various characters. This
agrees with Addisu Fekadu et al. (2013) who reported
higher genotypic correlation than phenotypic
correlation. The main genetic cause of such
correlation is pleiotropy, which refers to manifold
effects of a gene (Falconer, 1989).
Above results were similarly reported by Tripura et
al. (2016) who found tuber number per plant have
positive and significant association with total tuber
yield. Panigrahi et al. (2017) reported that total tuber
yield showed positive and significant correlation with
marketable tuber yield at both phenotypic as well as
at genotypic levels. Sattar et al. (2007) reported tuber
yield per plant was positively and significantly
correlated with number of tubers per plant, average
tuber weight and dry matter content of tuber. He also
showed the significant positive genotypic correlation
of average weight of tubers with number of tubers
per plant, yield of tuber per plant and tuber dry
matter percentages.

Manamno et al. East African Journal of Sciences Volume 15 (1) 1-16

Table 6. Genotypic (above diagonal) and phenotypic (below diagonal) correlation coefficients for different pairs of traits in potato.

Trait DE DF DMA SH SN TYP TNP AW MTN MY TY DM SC SMN LSN

DE 1 0.24* 0.14 -0.21 -0.4** -0.15 -0.26* 0.1 -0.24* -0.09 -0.09 0.07 0.07 -0.09 0.14

DF 0.21* 1 -0.07 0.1 -0.15 0.11 0.01 0.15 0.00 0.12 0.14 0.05 0.05 -0.014 0.23*

DMA 0.04 -0.03 1 0.02 0.02 -0.03 0.07 0.02 0.04 0.07* 0.11* -0.03 -0.03 0.09 0.07

SH -0.17* 0.07 -0.02 1 0.1 0.102 0.73** 0.36** 0.43** 0.67** 0.71 ** 0.41** 0.41** -0.30* 0.43**

SN -0.29* -0.13 -0.04 0.07 1 0.13 0.53** -0.36** 0.42** 0.08 0.1 0.04 0.04 0.19 -0.39**

TYP -0.12 0.09 0.02 0.69** 0.11 1 0.5** 0.49 ** 0.61** 0.91** 0.97** 0.54** 0.54** -0.4** 0.54**

TNP -0.2* -0.00 -0.02 0.35** 0.45** 0.5** 1 -0.35* 0.8** 0.39* 0.47** 0.38* 0.38* 0.23* -0.35*

AW 0.08 0.13 0.03 0.38** -0.3** 0.59** -0.36** 1 -0.03 0.35* 0.59** 0.25* 0.25* -0.58** 0.90**

MTN -0.23* -0.02 -0.23* 0.38** 0.34** 0.57** 0.71** -0.05 1 0.7** 0.64** 0.47** 0.47** -0.36* -0.01

MY -0.10 0.10 0.05* 0.61** 0.06 0.89** 0.41** 0.54** 0.67** 1 0.95** 0.52** 0.52** -0.56** 0.61**

TY -0.08 0.11 0.06* 0.68** 0.07 0.95** 0.49** 0.54** 0.54** 0.93** 1 0.56** 0.56** -0.40* 0.57**

DM -0.06 0.04 0.03* 0.31** 0.05 0.45** 0.3** 0.21* 0.38** 0.44** 0.45** 1 1.00** -0.21 0.18

SC -0.06 0.04 0.03* 0.31** 0.05 0.45** 0.3** 0.21* 0.38** 0.44** 0.45** 1.0** 1 -0.21 0.18

SMN 0.08 0.01 -0.04 -0.14 0.08 -0.23* 0.16* -0.39** -0.52** -0.4** -0.20* -0.21* -0.21* 1 -0.64**

LSN 0.1 0.21* 0.05 0.39** -0.35 0.51** -0.34** 0.87** -0.03 0.57** 0.53** 0.14 0.14 0.14 1

Note: *and **, significant at P<0.05 and P<0.01, respectively, DE = days to emergency, DF = days to flowering, DMA = days to maturity, SH = plant stem height, SN = main stem number, TYP =
tuber yield per plant, TNP=tuber number per plant, AW = average tuber weight, MTN = marketable tuber number, MY = marketable yield, UNMY = unmarketable yield, TY = total tuber yield, SMN
= very small tuber size percentage, LSN = large size tuber percentage, DM = tuber dry matter content (%) and SC = tuber starch content (%).

Manamno et al. Genetic Variability among Potato Progenies

4. Conclusions
Genetic variability is the base for the crop
improvement. The availability of more diverse
current materials obtained from local crossing
indicated the chance of getting desirable genes to
improve the potato crop. Hence, potato offspring
generated from hybridization of cultivated potato
varieties showed genetically variability in yield and its
related traits and morphological traits in the current
study. The high range and mean values of the
evaluated parental varieties and progenies also suggest
that the existence of enough variability correlations of
traits. Biparental crosses of Jalene with Aterababa and
Belete with Aterababa varieties produced more
promising/better genotypes than the crosses of Gera
and Shenkola, and standard check variety in yield and
yield related traits. So, the result of this study also
indicated the continuously producing more diverse
clones by local crossing can reduce dependency on
international potato center materials in the country.
Therefore, additional experiments will be carried out
to evaluate the most promising genotypes for
desirable traits, with the purpose to either produce
new variety or select parental lines for further
breeding.

5. Acknowledgments
We are indebted to Amhara Regional Agricultural
Research Institute (ARARI) and Adet Agricultural
Research Center for the financial support that
enabled us to do the research.

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