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Int. J. Agri. & Agri. R.
Missihoun et al.
Page 23
RESEARCH PAPER OPEN ACCESS
Phenotypic variability and racial classification of landraces of
sorghum (Sorghum bicolor (L.) Moench) collected in the
Northwestern Benin
Antoine Abel Missihoun1*
, Hubert Adoukonou-Sagbadja1
, Vincent Ezin2
, Paulin
Sedah1
, Rollande Aladé Dagba1
, Corneille Ahanhanzo1
, Clément Agbangla1
1
Department of Genetics and Biotechnology, University of Abomey-Calavi, Benin
2
Laboratoire de Biologie Végétale, Université d’Abomey-Calavi, Benin
Article published on October 12, 2015
Key words: Agro-morphological diversity, Botanical race, Quantitative descriptors, Qualitative descriptors,
Benin.
Abstract
A morpho-botanical analysis of 76 sorghum accessions collected from the department of Donga, the northwestern
part of Benin was carried out. An agro-morphological assessment of accessions was conducted based on 15
descriptors (10 quantitative traits and 5 qualitative traits). The experiment was laid at alpha lattice design with
three repetitions. Four races (guinea, durra, caudatum and bicolor) were identified with high proportion of
accessions of guinea race (86.84%) and low proportion of other races such as durra (5.26%), caudatum (2.63 %)
and bicolor (1.32%). Kafir race was missed from the collection. The analysis of agro-morphological characters of
accessions showed a significant phenotypic variability of sorghum local varieties kept and managed by farmers in
Donga. The principal component analysis (PCA) performed on quantitative traits showed three axes accounted
for nearly 70% of the total variation. The first axis represents plant height, diameter of third internodes, leaf
length and leaf width of the third internodes; the second axis accounts for length of peduncle while the third one
stands for the weight of 1,000 grains. On the basis of the quantitative and qualitative traits studied, ascending
hierarchical classification according to Ward aggregation criteria differentiated four groups structured around
key variables such as morpho-physiological characteristics of grains (color, size, degree of bitterness) and race.
* Corresponding Author: Antoine Abel Missihoun  missihoun_antoine@yahoo.fr
International Journal of Agronomy and Agricultural Research (IJAAR)
ISSN: 2223-7054 (Print) 2225-3610 (Online)
http://www.innspub.net
Vol. 7, No. 4, p. 23-35, 2015
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 24
Introduction
Sorghum bicolor (L.) Moench, is a monocotyledon
self-pollinating plant of tropical origin, belonging to
the family of Poaceae. It is cultivated in many tropical
regions of the world where it is used in human and
animal food and also in industries (Djè et al., 2007).
Among the economically important cereals in the
world, sorghum ranks fifth behind maize (Zea mays),
rice (Oryza spp.), wheat (Triticum spp.) and barley
(Hordeum spp.) (FAOSTAT, 2010). In the Sahelian
and semi-arid zones of Africa, sorghum is one of the
main resources of food (Djè et al., 2007). Its grains
are rich in carbohydrates and are appreciated for
their high energy value.
In Benin, sorghum is a staple food and ranks second
behind maize with an annual production of 168,090
tons in 2010 (FAOSTAT, 2010). Sorghum still
remains a traditional and ancient crop in the
northwest of the country despise its rank. Apart from
its use as food, some landraces of sorghum are grown
for their red pigment in the leaf sheaths and
sometimes in adjacent parts of stem: tinctorial
sorghum and coloring sorghum (Kayodé et al., 2005)
and others for their use in traditional medicine (for
instance the bitter yellow sorghum) (Kayodé et al.,
2005; Missihoun et al., 2012a).
The domestication of sorghum would probably began
in the Northeast Africa in a region located between
present-day Sudan and Ethiopia for about eight
thousand years ago (Wendorf et al., 1992). Sorghum
domesticated shows high morphological diversity in
terms of color and size of grains, color and length of
glumes, length and shape of panicles, and vegetative
cycle, etc. (Kayodé et al., 2006; Barnaud, 2007 ; Djè
et al., 2007; Barro-Kondombo et al., 2010), which
contributes to its remarkable adaptation to varying
climatic, soil and ecological conditions. The cultivated
sorghums are generally classified botanically and
based on the shape of the spikelet, grains and glumes
in 5 basic races (guinea, caudatum, durra, kafir,
bicolor) and 10 intermediate races gathering the
characteristics of two or more basic races (Harlan and
de Wet, 1972).
For better management, conservation and
valorization through genetic improvement of its
resources, the morphological diversity of cultivated
sorghum was assessed worldwide. This evaluation
prerequisite for any molecular genetics
characterization allows to detect possible synonyms
or homonyms of farmers’ classification of varieties
and also to make a reasoned and representative
choice of samples for analysis at molecular level. In
Africa, many studies of morphological diversity were
carried out. In Ethiopia, a study on 10 qualitative
traits and 54 fields showed the existence of significant
morphological variability between fields and that
observed from the enzyme data (Djè et al., 1998).
Many other studies assessing agro-morphological
variability of landraces of cultivated sorghum were
conducted. In West Africa, for instance, studies of
genetic diversity of local varieties of cultivated
sorghum were carried out by several researchers
(Morden et al., 1989; Zongo, 1991; Ollitrault et al.,
1997). Recently in Burkina Faso, a study of
phenotypic variability of local sorghum varieties in
two regions was performed (Barro-Kondombo et al.,
2008). It appears from this study that the racial
diversity of cultivated sorghum in these regions is low
and largely dominated by the guinea race
representing 96.8% of cultivated varieties. This study
confirmed that of Zongo (1991) also carried out in
Burkina Faso by taking into consideration the whole
country and estimated the proportion of accessions of
guinea race at 93.1%. This preponderance of guinea
race is not observed in other countries of the sub-
region such as Nigeria (40.2% of guinea), Niger
(10.5% of guinea) and Chad (7.0% of guinea) (Barro-
Kondombo et al., 2008, Deu et al., 2008). In Benin,
the agro-morphological characterization of sorghum
local varieties remains in small portions and data on
racial classification of cultivated accessions are almost
non-existent. However, the preservation of genetic
resources of cultivated species requires a thorough
knowledge of their phenotypic and genetic diversity,
for the purpose of better management and
valorization through plant improvement.
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 25
The present work aims at:
(1) Assessing agro-morphological characteristics of
accessions of cultivated sorghum in the northwestern
parts of Benin;
(2) Identifying botanically different races of cultivated
sorghum in the Northwestern Benin.
Materials and methods
Plant materials
The study was conducted on 76 accessions of local
varieties (Table 1) surveyed in December 2010 in
thirteen villages of Donga, the northwestern Benin
(Missihoun et al., 2012a) according to participatory
research method described by Tuan et al. (2003) and
Adoukonou-Sagbadja et al. (2006). Accessions were
local varieties grown by farmers from different ethnic
groups in the country.
Experimental site
The agro-morphological evaluation of accessions was
conducted at the experimental site on the campus of
University of Abomey-Calavi (6°24'57"N 2°20'31"E)
in Benin. It is a tropical wet zone characterized by
strong heat and high relative humidity ranging from
85-90%. The soil is ferralitic type dominated by
sandy-clayey sediments of the continental terminal.
Its pH is close to 6.2 and the organic matter content is
1.58%. The climate is of sub-equatorial type
characterized by a rainfall of 800 mm to 1200 mm
with four seasons: two rainy and two dry. The main
rainy season is from April to July and early
September to October. As for the dry season, the long
one is from November to March and the short one
from July to August (Table 2). The experiments were
conducted from May 2011 to January 2012.
Table 1. Code of the accessions, vernacular name, racial group and some information’s of grain and growing
season.
N°
Code of
the
Accessions
Vernacular
name
Racial
group
Colour of
grain
Colour of
husk
Panicle
shape
Growing
season
1 SOD 1 Zomouara G Red Black Loose Early-maturing
2 SOD 2 Zowémoha G Red Black Loose Early-maturing
3 SOD 3 Zomooha G Red Black Loose Early-maturing
4 SOD 4 Ano Eranan chinchin G Red Red Loose Early-maturing
5 SOD 5 Moussema G Red Black Loose Early-maturing
6 SOD 6 Zoténtérém DK Red Red Compact Early-maturing
7 SOD 7 Zomoaha G Red Black Loose Early-maturing
8 SOD 8 Zomooha 1 G Red Black Loose Early-maturing
9 SOD 9 Zomooha 2 G Red Purple Loose Early-maturing
10 SOD 10 M'ssema G Red Black Loose Early-maturing
11 SOD 11 Koussem G Red Black Loose Early-maturing
12 SOD 12 Zomoaha 1 G Red Black Loose Early-maturing
13 SOD 13 Anoro Kin'ka G Red Black Loose Early-maturing
14 SOD 14 Zoténtérém DK Red Red Compact Early-maturing
15 SOD 15 Zoumbouara G Red Black Loose Early-maturing
16 SOD 16 Aka Kpankpan Kékéré G Red Black Loose Early-maturing
17 SOD 17 Zopéra G White Black Loose Late-maturing
18 SOD 18 Zopira G White Black Loose Late-maturing
19 SOD 19 Zobeumdjoura rouge
(à glumes rouges)
G White Red Loose Late-maturing
20 SOD 20 Kèmnin Piha G White Black Loose Early-maturing
21 SOD 21 Agbani blanc G White Black Loose Late-maturing
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 26
N°
Code of
the
Accessions
Vernacular
name
Racial
group
Colour of
grain
Colour of
husk
Panicle
shape
Growing
season
22 SOD 22 Zobomdjouha G White Black Compact Late-maturing
23 SOD 23 Agbani 1 G White Red Loose Late-maturing
24 SOD 24 Tiam'la (blanc) G White Black Loose Early-maturing
25 SOD 25 Zopiha 1 A G White Black Loose Late-maturing
26 SOD 26 Zopiha 1 B KG White Black Compact Late-maturing
27 SOD 27 Zopiha 2 G White Black Loose Late-maturing
28 SOD 28 Ano Efoonon G White Black Loose Late-maturing
29 SOD 29 Koulom A G White Purple Loose Late-maturing
30 SOD 30 Koulom B G White Purple Loose Late-maturing
31 SOD 31 Aka Inodjo G Whitish Whitish Loose Early-maturing
32 SOD 32 Aka foufou Kékéré G White Black Loose Early-maturing
33 SOD 33 Agbani 2 G White Noirâtre Loose Late-maturing
34 SOD 34 Kouhloumè G White Red Loose Late-maturing
35 SOD 35 Aka foufou Lako G Whitish Black Loose Late-maturing
36 SOD 36 Sèmoutchè G White blackish Very
loose
Early-maturing
37 SOD 37 Anoro Foonon 1 G White
speckled with
crimson
blackish Loose Late-maturing
38 SOD 38 Anoro Foonon 2 G Whitish blackish Loose Late-maturing
39 SOD 39 Vèmah 1 DG White Purple Compact Early-maturing
40 SOD 40 Zomoala D Yellow White Compact Intermediate-
maturing
41 SOD 41 Ano chochomi D Yellow White Compact Intermediate-
maturing
42 SOD 42 Zomoila D Yellow White Compact Intermediate-
maturing
43 SOD 43 Zomoora D Yellow White Compact Intermediate-
maturing
44 SOD 44 Zotihou D Yellow White Compact Intermediate-
maturing
45 SOD 45 Sèmgin moaha G Whitish Red/Black Loose Intermediate-
maturing
46 SOD 46 Lam'za moaha G Whitish Red/Black Loose Intermediate-
maturing
47 SOD 47 Zogawa G Whitish Crimson Loose Intermediate-
maturing
48 SOD 48 Narabeunzo B blackish Black Loose -
49 SOD 49 Zokaram nini 1 G Reddish Red Loose Late-maturing
50 SOD 50 Zokaram nini 2 G Whitish Black Loose Late-maturing
51 SOD 51 Lam néza G Reddish/
Whitish
Black Loose
52 SOD 52 Lamza G Whitish Black Loose Intermediate-
maturing
53 SOD 53 Sèmgnin Piha G Whitish White-
Black
Loose Intermediate-
maturing
54 SOD 54 Ano Kaki G Reddish Black Loose Early-maturing
55 SOD 55 Ano Eranan G Reddish Red Loose Intermediate-
maturing
56 SOD 56 Za (moins rouge) G Whitish Red Loose Late-maturing
57 SOD 57 Agbani (rouge) G Whitish Crimson Loose Late-maturing
58 SOD 58 Za (rouge) ou Zaga G Whitish Red Loose Late-maturing
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 27
N°
Code of
the
Accessions
Vernacular
name
Racial
group
Colour of
grain
Colour of
husk
Panicle
shape
Growing
season
59 SOD 59 B'mdjoura G Whitish Red Loose Late-maturing
60 SOD 60 Zokpénaï D Yellowish White Compact Early-maturing
61 SOD 61 Zooga G White/Red Black Loose Late-maturing
62 SOD 62 Talèm'la G Whitish Red Loose Late-maturing
63 SOD 63 Zoniniléti G Whitish Black Loose Late-maturing
64 SOD 64 M'la Kssèmèè G Reddish Red Loose Early-maturing
65 SOD 65 Anoro Kin'ka 2 G Reddish Red Loose Intermediate-
maturing
66 SOD 66 Moukoulikouté C Reddish/
Yellowish
Whitish Compact Late-maturing
67 SOD 67 Zobeundjouha G White/Red Red Semi-
loose
Late-maturing
68 SOD 68 Moussii D Yellowish White Compacte Early-maturing
69 SOD 69 Zomoaha 2 G Reddish Red Loose Early-maturing
70 SOD 70 M'sséé C Yellowish Crimson Compact Early-maturing
71 SOD 71 Vèmah 2 D Whitish Yellowish Compact Early-maturing
72 SOD 72 Talèm'la G Whitish Black Loose Late-maturing
73 SOD 73 Zoumboua G Reddish Black Loose Early-maturing
74 SOD 74 aka kpankpan Lako 1 G Reddish Red Loose Early-maturing
75 SOD 75 aka kpankpan Lako 2 G Whitish Reddish Loose Early-maturing
76 SOD 76 Zowémoha 2 G Reddish Black Loose Early-maturing
Table 2. Climatic characteristics of the sampled villages and experimental site.
Townships of
department
Villages sampled
Climatic characteristics (adapted
from WorldClim data base)
Other crops grown in
addition to sorghum
Number
of samples
Bassila
Guiguissou
Salmanga
Climate : Sudano-Guinean
rainfall: 1150-1183 mm with
unimodal regime (May-October)
Temperature: 20°C-33°C
Millet, maize, yam,
groundnut, cassava,
cowpea, pepper,
tomato, okra, cashew
9
Patargo
Climate : Sudanese
rainfall : 1250 mm on average with
unimodal regime (May-September)
Temperature: 20°C-33°C
Millet, maize, yam,
groundnut, cassava,
pepper, tomato, okra,
rice, cashew
6
Djougou
Danogou Onklou
Darawinga
Pélébina Serou
Yoroussonga
Climate : Sudanese
rainfall : 1130-1164 mm with
unimodal regime (May-
September)
Temperature: 20°C-33°C
Millet, yam, rice,
maize, groundnut,
cassava, potato,
cowpea, soya, bambara
groundnut, pepper,
tomato, okra, cashew
37
Copargo
Kpassabega
Singre
Climate : Sudanese
Rainfall : 1153-1164 mm with
unimodal regime (May-
September)
Temperature: 20°C-33°C
Millet, yam, rice,
maize, groundnut,
cassava, potato,
cowpea, tomato, okra,
cashew
16
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 28
Townships of
department
Villages sampled
Climatic characteristics (adapted
from WorldClim data base)
Other crops grown in
addition to sorghum
Number
of samples
Ouaké
Kim-Kim
Sonaholou
Climate: Sudanese
Rainfall : 1227-1262 mm with
unimodal regime (May-
September)
Temperature: 20°C-33°C
Millet, maize, yam,
groundnut, cassava,
cowpea, rice, bambara
groundnut, pepper,
tomato, okra, cashew
8
Abomey-
Calavi
(experimental
site)
University of
Abomey-Calavi
(experimental
site)
Climate : Guinean (sub-equatorial
type)
rainfall : 1160 mm on average with
bimodal regime (April-July and
from September-October)
Temperature: 24°C-31°C
Non documented
76 (Total
of
samples)
Experimental design
The experimental design was laid at alpha lattice
(Patterson and Williams, 1976) with three replicates,
11 blocks and 11 accessions per entry. Each accession
was randomly sown in the three replicates. For each
accession, the plot unit was 1.6 m long and 1 m wide
with 0.80 m line spacing and 0.20 m row spacing.
Each plot unit consisted of 10 plants. Only one plant
was left in seed hole 10 days after emergence (Fig. 1a).
The parameters were measured on five plants
randomly selected from each primary plot. These
plants were monitored until maturity (Fig. 1b and Fig.
1c) and various quantitative and qualitative
parameters were recorded.
A total of 5 quantitative parameters and 5 qualitative
parameters were measured during the
experimentation. Quantitative parameters measured
were: plant height (Hpl), length of the panicle (Lpa),
peduncle length (Lpe), diameter of the third
internodes (Dte), leaf length of the third internodes
(Ltf ), leaf width of the third internodes (LTF), the
panicle exsertion (exp), number of leaves (NFE), the
effective number of tillers (Nw) and the weight of
1000 grains (p ~ 1000). The qualitative parameters
measured were the emergence vigor (Vem), grain
color (Cgr), color of glumes (Cgl), panicle type (Ptp)
and grain type (GTO). The various parameters were
recorded according to the recommendations of the
revised descriptors of sorghum (Bioversity
International and ICRISAT, 1993).
As for racial classification, it was done according to
the criteria defined by Harlan and de Wet (1972) in
two stages: initially on panicles collected from
farmers and secondly on panicles harvested in the
end of the experiment carried out at the Laboratory of
Genetics and Biotechnology, University of Abomey-
Calavi.
Data analysis
In order to estimate the heterogeneity between the
blocks, uni-variate statistical analysis of variance
(ANOVA) with one factor was used with the aid of
Statistica 7.1 software. A principal component
analysis (PCA) was performed in order to estimate the
overall diversity within the studied population and to
describe the relationships between variables. This
analysis was followed by a hierarchical cluster
analysis (HCA). The grouping of accessions in
phenotypic classes by the method of hierarchical
cluster analysis was performed. From these results,
the number of classes and the class of each
composition were determined. Given that variables
are coded, descriptive statistics of the classes in the
table of frequencies allowed describing each of these
classes.
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 29
Fig. 1. Experimental site and some observations.
A: Overview of the experimental site after thinning to one plant per seed hole
B: early accessions with early flowering
C: Panicles of some accessions at maturity stage Traits measured
Results
Quantitative variation among accessions analyzed
The overall variability of traits is presented in Table 3.
Plant height varies from 213.33 cm to 486.36 cm
while the length of the panicle ranges from 17.63 cm
to 48.60 cm. Peduncle length ranges from 18.00 cm
to 78.50 cm and the diameter of the third internodes
from 0.55 cm to 1.30 cm. As for the leaf length and
leaf width of the third internodes they vary from 41.85
cm to 98.89 cm and from 3.55 cm to 8.83 cm
respectively while the panicle exsertion ranges from
0.00 cm to 34.37 cm. The number of leaves per plant
varies from 16 cm to 28 cm while the number of
effective tillers was 1 or 2. The weight of 1000 grains
accessions ranges from 7.50 g to 52.80 g. The
variables with significant variation were panicle
length, peduncle length, leaf length of the third
internodes, panicle exsertion and weight of 1000
grains (Fig. 2).
Table 3. Performance of genotypes of sorghum
accessions (Sorghum bicolor (L.) Moench) analyzed.
Quantitative
traits
Mini-
mum
Maxi-
mum
Average
Standard
deviation
Hpl (cm) 213.33 486.36 339.57 60.83
Lpa (cm) 17.63 48.60 33.43 6.83
Lpe (cm) 18.00 78.50 43.89 11.17
Dte (cm) 0.55 1.30 0.93 0.16
Ltf (cm) 41.85 98.89 74.82 9.91
ltf (cm) 3.55 8.83 5.59 1.05
Exp (cm) 0.00 34.37 10.08 6.59
A
C
B
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 30
Quantitative
traits
Mini-
mum
Maxi-
mum
Average
Standard
deviation
Nfe (Number) 16.00 27.67 20.47 2.35
Nte (Number) 1.00 1.75 1.08 0.17
p~1000 (g) 7.50 52.80 32.63 8.07
Hpl = plant height, Lpa = panicle length, Lpe = peduncle
length, Dte = diameter of the third internodes, Ltf = leaf
length of the third internodes, ltf = leaf width of the third
internodes, Exp = panicle exsertion, Nfe = number of
leaves, Nw = number of effective tillers and p = ~ 1000g
weight of 1,000 grains.
Fig. 2. Global Change in quantitative variables within
accessions.
Correlation between quantitative variables
The correlation matrix between the measured
variables is presented in Table 4. The analysis of this
table shows that plant height was significantly and
positively correlated with other variables except for
panicle exsertion and number of effective tillers. The
panicle length was positively correlated with the
length of the peduncle, the leaf length and leaf width
of the third internodes and the weight of 1000 grain
(p ~ 1000). Peduncle length was negatively correlated
to the diameter of the third internodes and number of
leaves but was significantly and positively correlated
with panicle exsertion.
The diameter of the third internodes was positively
correlated with the leaf length and leaf width of the
third internodes and the number of leaves but
negatively correlated to the panicle exsertion. Leaf
length of the third internodes was positively
correlated with leaf width, number of leaves and
weight of 1000 grains (p ~ 1000) and negatively
correlated with panicle exsertion. Leaf width of the
third internodes was positively correlated only to the
number of leaves. With regard to the panicle
exsertion and the effective number of tillers, they
were negatively correlated to the number of leaves
and the weight of 1000 grains (p ~ 1000) respectively
and the positively correlated to the weight of 1000
grains (p ~ 1000).
Table 4. Correlation matrix of quantitative variables measured.
Hpl Lpa Lpe Dte Ltf ltf Exp Nfe Nte p~1000
Hpl 1.00
Lpa 0.73* 1.00
Lpe 0.31* 0.28* 1.00
Dte 0.41* 0.20 -0.28* 1.00
Ltf 0.53* 0.45* -0.18 0.50* 1.00
ltf 0.43* 0.23* -0.11 0.73* 0.48* 1.00
Exp -0.02 -0.15 0.74* -0.24* -0.39* -0.01 1.00
Nfe 0.28* 0.09 -0.37* 0.44* 0.39* 0.47* -0.27* 1.00
Nte -0.12 -0.14 -0.07 -0.02 0.01 -0.02 -0.02 0.07 1.00
p~1000 0.34* 0.25* -0.11 0.13 0.43* -0.01 -0.32* 0.10 0.24* 1.00
* Significant correlation at 5%.
PCA analysis revealed variations within sorghum
landraces
The principal component analysis (PCA) was used to
explain the quantitative variations observed within
the plant material used. The correlations between
variables and the synthetic axes are showed in Fig. 3.
Axes 1 and 2 put together account for 54.91% of the
total variability in the sorghum accessions studied
(Fig. 3). As for the factorial scheme (axis 1 and axis 3),
it stands for 48.50% of the total variability. Overall,
the first three axes account for nearly 70% of the total
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 31
variation. These three axes were used to describe the
variability within the collection.
Fig. 3. Circle of correlation (ACP) in the plane 1-2.
Plant height, panicle length, diameter of the third
internodes, length and width of the third leaf and the
number of leaves were significantly and positively
correlated with axis 1. On the other hand, axis 2 was
significantly and negatively correlated with the length
of the peduncle and panicle exsertion. As for the
weight of 1000 grains, this variable was significantly
and negatively correlated to the main axis 3 (Table 5).
These variables were analyzed and interpreted
respectively in accordance with their respective axis.
Table 5. Coordinates of variable on the first three
axes.
Variables axis 1 axis 2 axis 3
Hpl 0.700796* -0.595646 -0.122998
Lpa 0.559952 -0.562835 -0.312229
Lpe -0.278482 -0.904343* -0.017289
Dte 0.762000* 0.091818 0.397449
Ltf 0.826384* -0.024270 -0.168076
Ltf 0.702494* -0.091277 0.555841
Exp -0.464255 -0.642451 0.366395
Nfe 0.620137 0.283847 0.295721
Nte -0.003610 0.241112 -0.281302
p~1000 0.437383 0.049252 -0.714529*
Grouping accessions on the basis of quantitative and
qualitative variables
A hierarchical classification of sorghum accessions
studied and performed with the aggregation criteria
of Ward from adjusted mean values for 15 variables
(10 quantitative and qualitative 5) from analysis of
variance results in the dendrogram (Fig. 4). The
Analysis of this dendrogram was able to differentiate
four main groups by truncation at a distance of
aggregation of 19. Group I contains 13 accessions that
were mainly from durra race with bitter or no bitter
yellow grains (SOD 40, SOD 41, SOD 42, SOD 44 and
SOD 60), an accession with yellowish grains of
caudatum race (SOD 71) and an accession of coloring
sorghum of intermediate race durra-kafir (SOD 6).
Group II composes of 24 accessions which were
mainly sorghum accessions of guinea race with long
and white grains of which vegetative cycle was long
(SOD 21, SOD 22, SOD 33, SOD35, SOD 37, SOD 47,
SOD 57 and SOD 62) and also contains some
accessions with red grains as SOD 2, SOD 4 and SOD
65 (all belonging to guinea race). Group III consists of
20 accessions which were essentially guinea race with
white or whitish grains and with medium growth
cycle (SOD 24, SOD 29, SOD31, SOD 36, SOD54, SOD
56, SOD59, and SOD 61). The accession of the bicolor
race (SOD 48) and an accession of caudatum race
(SOD 66) were also found in this group. Finally,
group VI consists of 17 accessions, mainly early
guinea race with red grains of small size (SOD 7, SOD
10, SOD 13, SOD 15, SOD 16, and SOD 64) but also
some accessions with white grains (SOD 17, SOD 26
and SOD 38).
Fig. 4. Ascending hierarchical classification (AHC) of
74 sorghum accessions according to the Ward
aggregation criteria.
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 32
Racial classification of sorghum accessions
76 sorghum accessions collected belong to four
botanical races (guinea, Durra, caudatum and bicolor)
out of the existing five races. The guinea race was
predominant and represents 86.84% of the collection
followed by durra race with 5.26%, caudatum race
with 2.63% and bicolor race with 1.32%. The
intermediaries represent 3.95% (Fig. 5, Table 1). In
terms of grain color, 41.33% had an intermediate
color (pink, reddish, whitish, yellowish, blackish,
white-spotted purple or red, multicolored, etc.),
30.66% of the grains were white, 21.33% were red and
6.66% were yellow. Moreover, it should be noted that
53.95% of accessions had black glumes, 80.26% had
loose panicles and 42.11% had a short vegetative
cycle.
Fig. 5. Proportional distribution of different
botanical races of accessions analyzed.
Discussion
Multivariate analysis of quantitative and qualitative
data revealed a structuring in 4 distinct groups of
sorghum accessions based mainly on racial factor, but
also the duration of the vegetative cycle and morpho-
physiological characteristics of grains (color, size,
degree of bitterness). This structuring of Benin
accessions according to botanical race confirms once
again previous agro-morphological investigations on
collections from countries of West Africa (Deu et al.,
2006; Barro-Kondombo et al., 2008.). The
structuring according to morpho-physiological
characteristics of the grain was also reported in some
studies such as those conducted by Barro-Kondombo
et al. (2008) who identified, after agro-morphological
characterization, an original group of accessions of
local varieties of sorghum with red grains. In the
present study, the group of accessions that appears
more homogeneous on the basis of grain color is
group I which consists of accessions with bitter or no
bitter yellow grains. This phenotypic difference of this
group of accessions appears extremely important for
the judicious use in research because sorghum local
varieties with bitter yellow grains are widely
recommended for use in traditional medicine in
Benin (Kayodé et al., 2005; Missihoun et al., 2012a).
In addition, groups II, III, and IV compose of
landraces of guinea race which differ in the duration
of vegetative cycle and the nature of the grains. These
two factors also seem to play an important role in
agro-morphological structuring of accessions of
cultivated sorghum in Burkina Faso (Barro-
Kondombo et al., 2008). However, in Benin, ethno-
botanical investigations revealed the specific
importance of each of these different groups of local
varieties. Local varieties with red grains and short
growth cycle (group IV) currently dominate Benin
sorghum due to their relative tolerance to drought,
their earliness (early maturity), their adaptability to
relatively poor soils and their high market value.
Local varieties with white grains or intermediate
color, medium or long growth cycle (group II and
group III) are often more profitable in terms of
production of grains, more farinaceous and are used
in the production of local dough (Wô, Dibou, Sifanou,
Foura). They are sensitive to drought and require soil
fertility. Finally, local varieties bitter yellow grains
(group I) were mainly grown for use in traditional
medicine. This structuring observed with the agro-
morphological traits was globally confirmed by
molecular analyzes with nuclear microsatellite
markers used on the same collection (Missihoun et
al., 2015).
In terms of selection and/or varietal improvement,
positive correlations observed between the different
quantitative variables showed that the accessions with
long panicles were of big size (big plant height) and
those with the weight of 1000 grains had the largest
Int. J. Agri. & Agri. R.
Missihoun et al.
Page 33
diameter of the third internodes. On-farm, farmers
had a preference for varieties of high size due to both
the quality of their food grains and especially to their
important role they play in the production of biomass
for combustion and building materials (Kayodé et al.,
2006). The utilization of these results will be very
important in plant breeding programs to improve the
performance of local varieties in terms of grain and
biomass production.
The racial diversity of cultivated sorghum in Donga
department is very high (presence of 4 races out of
the five existing races) with a predominance of guinea
race. The dominant presence of guinea race in the
regions of West Africa is known recognized and has
been reported by many researchers from West African
countries such as Burkina Faso (Ollitrault 1987;
Zongo, 1991; Barro-Kondombo et al., 2010), Mali
(Sagnard et al., 2008), Ghana (Kudadjie, 2006) and
even in Central Africa with Cameroon (Barnaud,
2007). The present study is consistence with those
reported above. The results of this work in Benin
could be justified in terms of the geographical
position of the country in relation to the history of
diversification of sorghum. As a matter of fact, Benin
is located in the central of West African countries
where there is the diversification of guinea race and
with high likelihood of plant materials (seeds)
exchanges between farmers of the neighboring
countries such as Burkina Faso and Niger. The
predominance of guinea race could also be explained
by climatic factor. Guinea race would be more suited
to the higher rainfall areas (Barro-Kondombo et al.,
2010). Furthermore, the absence of kafir race among
collected samples in Benin confirms all the previous
work done in the regions of West Africa (Burkina
Faso, Niger, and Ghana). Deu et al. (2008) observed
the same racial diversity in Niger (all races except
Kafir) with the difference that their proportion were
not the same: durra (23.1%), caudatum (21.3%),
guinea (10.5%) and bicolor (8.1%). Kafir race is
reported to be found in southern Africa and is almost
absent in the regions of West Africa, although more
intermediate of this race with other races exist.
Conclusion
This study is one of the first evaluations which
focused on agro-morphological characteristics of
grown sorghum showed an important phenotypic
variability of local varieties of sorghum kept and
managed by farmers in Donga. Based on the traits
studied four groups structured around factors such as
morpho-physiological characteristics of the grains
(color, size, taste) and race, were identified.
Moreover, the results showed the presence of four
pure races (guinea, Durra, caudatum bicolor) and the
intermediaries such as guinea-durra, guinea-kafir,
kafir-durra in the surveyed areas. Guinea race is
predominant among the races. The use of molecular
genetic markers in further work will allow thorough
analysis of the evolution factors of genetic diversity
for a better use of local resources of grown sorghum
in breeding and conservation programs. The interest
in the long run, is the sustainability of sorghum that
will continue to play its role in food security and the
diversification of cereal production country.
Acknowledgment
The authors thank Mrs Avohou Germaine for her help
for collecting data. We thank also the anonymous
reviewers for their valuable comments.
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Phenotypic variability and racial classification of landraces of sorghum (Sorghum bicolor (L.) Moench) collected in the Northwestern Benin

  • 1. Int. J. Agri. & Agri. R. Missihoun et al. Page 23 RESEARCH PAPER OPEN ACCESS Phenotypic variability and racial classification of landraces of sorghum (Sorghum bicolor (L.) Moench) collected in the Northwestern Benin Antoine Abel Missihoun1* , Hubert Adoukonou-Sagbadja1 , Vincent Ezin2 , Paulin Sedah1 , Rollande Aladé Dagba1 , Corneille Ahanhanzo1 , Clément Agbangla1 1 Department of Genetics and Biotechnology, University of Abomey-Calavi, Benin 2 Laboratoire de Biologie Végétale, Université d’Abomey-Calavi, Benin Article published on October 12, 2015 Key words: Agro-morphological diversity, Botanical race, Quantitative descriptors, Qualitative descriptors, Benin. Abstract A morpho-botanical analysis of 76 sorghum accessions collected from the department of Donga, the northwestern part of Benin was carried out. An agro-morphological assessment of accessions was conducted based on 15 descriptors (10 quantitative traits and 5 qualitative traits). The experiment was laid at alpha lattice design with three repetitions. Four races (guinea, durra, caudatum and bicolor) were identified with high proportion of accessions of guinea race (86.84%) and low proportion of other races such as durra (5.26%), caudatum (2.63 %) and bicolor (1.32%). Kafir race was missed from the collection. The analysis of agro-morphological characters of accessions showed a significant phenotypic variability of sorghum local varieties kept and managed by farmers in Donga. The principal component analysis (PCA) performed on quantitative traits showed three axes accounted for nearly 70% of the total variation. The first axis represents plant height, diameter of third internodes, leaf length and leaf width of the third internodes; the second axis accounts for length of peduncle while the third one stands for the weight of 1,000 grains. On the basis of the quantitative and qualitative traits studied, ascending hierarchical classification according to Ward aggregation criteria differentiated four groups structured around key variables such as morpho-physiological characteristics of grains (color, size, degree of bitterness) and race. * Corresponding Author: Antoine Abel Missihoun  missihoun_antoine@yahoo.fr International Journal of Agronomy and Agricultural Research (IJAAR) ISSN: 2223-7054 (Print) 2225-3610 (Online) http://www.innspub.net Vol. 7, No. 4, p. 23-35, 2015
  • 2. Int. J. Agri. & Agri. R. Missihoun et al. Page 24 Introduction Sorghum bicolor (L.) Moench, is a monocotyledon self-pollinating plant of tropical origin, belonging to the family of Poaceae. It is cultivated in many tropical regions of the world where it is used in human and animal food and also in industries (Djè et al., 2007). Among the economically important cereals in the world, sorghum ranks fifth behind maize (Zea mays), rice (Oryza spp.), wheat (Triticum spp.) and barley (Hordeum spp.) (FAOSTAT, 2010). In the Sahelian and semi-arid zones of Africa, sorghum is one of the main resources of food (Djè et al., 2007). Its grains are rich in carbohydrates and are appreciated for their high energy value. In Benin, sorghum is a staple food and ranks second behind maize with an annual production of 168,090 tons in 2010 (FAOSTAT, 2010). Sorghum still remains a traditional and ancient crop in the northwest of the country despise its rank. Apart from its use as food, some landraces of sorghum are grown for their red pigment in the leaf sheaths and sometimes in adjacent parts of stem: tinctorial sorghum and coloring sorghum (Kayodé et al., 2005) and others for their use in traditional medicine (for instance the bitter yellow sorghum) (Kayodé et al., 2005; Missihoun et al., 2012a). The domestication of sorghum would probably began in the Northeast Africa in a region located between present-day Sudan and Ethiopia for about eight thousand years ago (Wendorf et al., 1992). Sorghum domesticated shows high morphological diversity in terms of color and size of grains, color and length of glumes, length and shape of panicles, and vegetative cycle, etc. (Kayodé et al., 2006; Barnaud, 2007 ; Djè et al., 2007; Barro-Kondombo et al., 2010), which contributes to its remarkable adaptation to varying climatic, soil and ecological conditions. The cultivated sorghums are generally classified botanically and based on the shape of the spikelet, grains and glumes in 5 basic races (guinea, caudatum, durra, kafir, bicolor) and 10 intermediate races gathering the characteristics of two or more basic races (Harlan and de Wet, 1972). For better management, conservation and valorization through genetic improvement of its resources, the morphological diversity of cultivated sorghum was assessed worldwide. This evaluation prerequisite for any molecular genetics characterization allows to detect possible synonyms or homonyms of farmers’ classification of varieties and also to make a reasoned and representative choice of samples for analysis at molecular level. In Africa, many studies of morphological diversity were carried out. In Ethiopia, a study on 10 qualitative traits and 54 fields showed the existence of significant morphological variability between fields and that observed from the enzyme data (Djè et al., 1998). Many other studies assessing agro-morphological variability of landraces of cultivated sorghum were conducted. In West Africa, for instance, studies of genetic diversity of local varieties of cultivated sorghum were carried out by several researchers (Morden et al., 1989; Zongo, 1991; Ollitrault et al., 1997). Recently in Burkina Faso, a study of phenotypic variability of local sorghum varieties in two regions was performed (Barro-Kondombo et al., 2008). It appears from this study that the racial diversity of cultivated sorghum in these regions is low and largely dominated by the guinea race representing 96.8% of cultivated varieties. This study confirmed that of Zongo (1991) also carried out in Burkina Faso by taking into consideration the whole country and estimated the proportion of accessions of guinea race at 93.1%. This preponderance of guinea race is not observed in other countries of the sub- region such as Nigeria (40.2% of guinea), Niger (10.5% of guinea) and Chad (7.0% of guinea) (Barro- Kondombo et al., 2008, Deu et al., 2008). In Benin, the agro-morphological characterization of sorghum local varieties remains in small portions and data on racial classification of cultivated accessions are almost non-existent. However, the preservation of genetic resources of cultivated species requires a thorough knowledge of their phenotypic and genetic diversity, for the purpose of better management and valorization through plant improvement.
  • 3. Int. J. Agri. & Agri. R. Missihoun et al. Page 25 The present work aims at: (1) Assessing agro-morphological characteristics of accessions of cultivated sorghum in the northwestern parts of Benin; (2) Identifying botanically different races of cultivated sorghum in the Northwestern Benin. Materials and methods Plant materials The study was conducted on 76 accessions of local varieties (Table 1) surveyed in December 2010 in thirteen villages of Donga, the northwestern Benin (Missihoun et al., 2012a) according to participatory research method described by Tuan et al. (2003) and Adoukonou-Sagbadja et al. (2006). Accessions were local varieties grown by farmers from different ethnic groups in the country. Experimental site The agro-morphological evaluation of accessions was conducted at the experimental site on the campus of University of Abomey-Calavi (6°24'57"N 2°20'31"E) in Benin. It is a tropical wet zone characterized by strong heat and high relative humidity ranging from 85-90%. The soil is ferralitic type dominated by sandy-clayey sediments of the continental terminal. Its pH is close to 6.2 and the organic matter content is 1.58%. The climate is of sub-equatorial type characterized by a rainfall of 800 mm to 1200 mm with four seasons: two rainy and two dry. The main rainy season is from April to July and early September to October. As for the dry season, the long one is from November to March and the short one from July to August (Table 2). The experiments were conducted from May 2011 to January 2012. Table 1. Code of the accessions, vernacular name, racial group and some information’s of grain and growing season. N° Code of the Accessions Vernacular name Racial group Colour of grain Colour of husk Panicle shape Growing season 1 SOD 1 Zomouara G Red Black Loose Early-maturing 2 SOD 2 Zowémoha G Red Black Loose Early-maturing 3 SOD 3 Zomooha G Red Black Loose Early-maturing 4 SOD 4 Ano Eranan chinchin G Red Red Loose Early-maturing 5 SOD 5 Moussema G Red Black Loose Early-maturing 6 SOD 6 Zoténtérém DK Red Red Compact Early-maturing 7 SOD 7 Zomoaha G Red Black Loose Early-maturing 8 SOD 8 Zomooha 1 G Red Black Loose Early-maturing 9 SOD 9 Zomooha 2 G Red Purple Loose Early-maturing 10 SOD 10 M'ssema G Red Black Loose Early-maturing 11 SOD 11 Koussem G Red Black Loose Early-maturing 12 SOD 12 Zomoaha 1 G Red Black Loose Early-maturing 13 SOD 13 Anoro Kin'ka G Red Black Loose Early-maturing 14 SOD 14 Zoténtérém DK Red Red Compact Early-maturing 15 SOD 15 Zoumbouara G Red Black Loose Early-maturing 16 SOD 16 Aka Kpankpan Kékéré G Red Black Loose Early-maturing 17 SOD 17 Zopéra G White Black Loose Late-maturing 18 SOD 18 Zopira G White Black Loose Late-maturing 19 SOD 19 Zobeumdjoura rouge (à glumes rouges) G White Red Loose Late-maturing 20 SOD 20 Kèmnin Piha G White Black Loose Early-maturing 21 SOD 21 Agbani blanc G White Black Loose Late-maturing
  • 4. Int. J. Agri. & Agri. R. Missihoun et al. Page 26 N° Code of the Accessions Vernacular name Racial group Colour of grain Colour of husk Panicle shape Growing season 22 SOD 22 Zobomdjouha G White Black Compact Late-maturing 23 SOD 23 Agbani 1 G White Red Loose Late-maturing 24 SOD 24 Tiam'la (blanc) G White Black Loose Early-maturing 25 SOD 25 Zopiha 1 A G White Black Loose Late-maturing 26 SOD 26 Zopiha 1 B KG White Black Compact Late-maturing 27 SOD 27 Zopiha 2 G White Black Loose Late-maturing 28 SOD 28 Ano Efoonon G White Black Loose Late-maturing 29 SOD 29 Koulom A G White Purple Loose Late-maturing 30 SOD 30 Koulom B G White Purple Loose Late-maturing 31 SOD 31 Aka Inodjo G Whitish Whitish Loose Early-maturing 32 SOD 32 Aka foufou Kékéré G White Black Loose Early-maturing 33 SOD 33 Agbani 2 G White Noirâtre Loose Late-maturing 34 SOD 34 Kouhloumè G White Red Loose Late-maturing 35 SOD 35 Aka foufou Lako G Whitish Black Loose Late-maturing 36 SOD 36 Sèmoutchè G White blackish Very loose Early-maturing 37 SOD 37 Anoro Foonon 1 G White speckled with crimson blackish Loose Late-maturing 38 SOD 38 Anoro Foonon 2 G Whitish blackish Loose Late-maturing 39 SOD 39 Vèmah 1 DG White Purple Compact Early-maturing 40 SOD 40 Zomoala D Yellow White Compact Intermediate- maturing 41 SOD 41 Ano chochomi D Yellow White Compact Intermediate- maturing 42 SOD 42 Zomoila D Yellow White Compact Intermediate- maturing 43 SOD 43 Zomoora D Yellow White Compact Intermediate- maturing 44 SOD 44 Zotihou D Yellow White Compact Intermediate- maturing 45 SOD 45 Sèmgin moaha G Whitish Red/Black Loose Intermediate- maturing 46 SOD 46 Lam'za moaha G Whitish Red/Black Loose Intermediate- maturing 47 SOD 47 Zogawa G Whitish Crimson Loose Intermediate- maturing 48 SOD 48 Narabeunzo B blackish Black Loose - 49 SOD 49 Zokaram nini 1 G Reddish Red Loose Late-maturing 50 SOD 50 Zokaram nini 2 G Whitish Black Loose Late-maturing 51 SOD 51 Lam néza G Reddish/ Whitish Black Loose 52 SOD 52 Lamza G Whitish Black Loose Intermediate- maturing 53 SOD 53 Sèmgnin Piha G Whitish White- Black Loose Intermediate- maturing 54 SOD 54 Ano Kaki G Reddish Black Loose Early-maturing 55 SOD 55 Ano Eranan G Reddish Red Loose Intermediate- maturing 56 SOD 56 Za (moins rouge) G Whitish Red Loose Late-maturing 57 SOD 57 Agbani (rouge) G Whitish Crimson Loose Late-maturing 58 SOD 58 Za (rouge) ou Zaga G Whitish Red Loose Late-maturing
  • 5. Int. J. Agri. & Agri. R. Missihoun et al. Page 27 N° Code of the Accessions Vernacular name Racial group Colour of grain Colour of husk Panicle shape Growing season 59 SOD 59 B'mdjoura G Whitish Red Loose Late-maturing 60 SOD 60 Zokpénaï D Yellowish White Compact Early-maturing 61 SOD 61 Zooga G White/Red Black Loose Late-maturing 62 SOD 62 Talèm'la G Whitish Red Loose Late-maturing 63 SOD 63 Zoniniléti G Whitish Black Loose Late-maturing 64 SOD 64 M'la Kssèmèè G Reddish Red Loose Early-maturing 65 SOD 65 Anoro Kin'ka 2 G Reddish Red Loose Intermediate- maturing 66 SOD 66 Moukoulikouté C Reddish/ Yellowish Whitish Compact Late-maturing 67 SOD 67 Zobeundjouha G White/Red Red Semi- loose Late-maturing 68 SOD 68 Moussii D Yellowish White Compacte Early-maturing 69 SOD 69 Zomoaha 2 G Reddish Red Loose Early-maturing 70 SOD 70 M'sséé C Yellowish Crimson Compact Early-maturing 71 SOD 71 Vèmah 2 D Whitish Yellowish Compact Early-maturing 72 SOD 72 Talèm'la G Whitish Black Loose Late-maturing 73 SOD 73 Zoumboua G Reddish Black Loose Early-maturing 74 SOD 74 aka kpankpan Lako 1 G Reddish Red Loose Early-maturing 75 SOD 75 aka kpankpan Lako 2 G Whitish Reddish Loose Early-maturing 76 SOD 76 Zowémoha 2 G Reddish Black Loose Early-maturing Table 2. Climatic characteristics of the sampled villages and experimental site. Townships of department Villages sampled Climatic characteristics (adapted from WorldClim data base) Other crops grown in addition to sorghum Number of samples Bassila Guiguissou Salmanga Climate : Sudano-Guinean rainfall: 1150-1183 mm with unimodal regime (May-October) Temperature: 20°C-33°C Millet, maize, yam, groundnut, cassava, cowpea, pepper, tomato, okra, cashew 9 Patargo Climate : Sudanese rainfall : 1250 mm on average with unimodal regime (May-September) Temperature: 20°C-33°C Millet, maize, yam, groundnut, cassava, pepper, tomato, okra, rice, cashew 6 Djougou Danogou Onklou Darawinga Pélébina Serou Yoroussonga Climate : Sudanese rainfall : 1130-1164 mm with unimodal regime (May- September) Temperature: 20°C-33°C Millet, yam, rice, maize, groundnut, cassava, potato, cowpea, soya, bambara groundnut, pepper, tomato, okra, cashew 37 Copargo Kpassabega Singre Climate : Sudanese Rainfall : 1153-1164 mm with unimodal regime (May- September) Temperature: 20°C-33°C Millet, yam, rice, maize, groundnut, cassava, potato, cowpea, tomato, okra, cashew 16
  • 6. Int. J. Agri. & Agri. R. Missihoun et al. Page 28 Townships of department Villages sampled Climatic characteristics (adapted from WorldClim data base) Other crops grown in addition to sorghum Number of samples Ouaké Kim-Kim Sonaholou Climate: Sudanese Rainfall : 1227-1262 mm with unimodal regime (May- September) Temperature: 20°C-33°C Millet, maize, yam, groundnut, cassava, cowpea, rice, bambara groundnut, pepper, tomato, okra, cashew 8 Abomey- Calavi (experimental site) University of Abomey-Calavi (experimental site) Climate : Guinean (sub-equatorial type) rainfall : 1160 mm on average with bimodal regime (April-July and from September-October) Temperature: 24°C-31°C Non documented 76 (Total of samples) Experimental design The experimental design was laid at alpha lattice (Patterson and Williams, 1976) with three replicates, 11 blocks and 11 accessions per entry. Each accession was randomly sown in the three replicates. For each accession, the plot unit was 1.6 m long and 1 m wide with 0.80 m line spacing and 0.20 m row spacing. Each plot unit consisted of 10 plants. Only one plant was left in seed hole 10 days after emergence (Fig. 1a). The parameters were measured on five plants randomly selected from each primary plot. These plants were monitored until maturity (Fig. 1b and Fig. 1c) and various quantitative and qualitative parameters were recorded. A total of 5 quantitative parameters and 5 qualitative parameters were measured during the experimentation. Quantitative parameters measured were: plant height (Hpl), length of the panicle (Lpa), peduncle length (Lpe), diameter of the third internodes (Dte), leaf length of the third internodes (Ltf ), leaf width of the third internodes (LTF), the panicle exsertion (exp), number of leaves (NFE), the effective number of tillers (Nw) and the weight of 1000 grains (p ~ 1000). The qualitative parameters measured were the emergence vigor (Vem), grain color (Cgr), color of glumes (Cgl), panicle type (Ptp) and grain type (GTO). The various parameters were recorded according to the recommendations of the revised descriptors of sorghum (Bioversity International and ICRISAT, 1993). As for racial classification, it was done according to the criteria defined by Harlan and de Wet (1972) in two stages: initially on panicles collected from farmers and secondly on panicles harvested in the end of the experiment carried out at the Laboratory of Genetics and Biotechnology, University of Abomey- Calavi. Data analysis In order to estimate the heterogeneity between the blocks, uni-variate statistical analysis of variance (ANOVA) with one factor was used with the aid of Statistica 7.1 software. A principal component analysis (PCA) was performed in order to estimate the overall diversity within the studied population and to describe the relationships between variables. This analysis was followed by a hierarchical cluster analysis (HCA). The grouping of accessions in phenotypic classes by the method of hierarchical cluster analysis was performed. From these results, the number of classes and the class of each composition were determined. Given that variables are coded, descriptive statistics of the classes in the table of frequencies allowed describing each of these classes.
  • 7. Int. J. Agri. & Agri. R. Missihoun et al. Page 29 Fig. 1. Experimental site and some observations. A: Overview of the experimental site after thinning to one plant per seed hole B: early accessions with early flowering C: Panicles of some accessions at maturity stage Traits measured Results Quantitative variation among accessions analyzed The overall variability of traits is presented in Table 3. Plant height varies from 213.33 cm to 486.36 cm while the length of the panicle ranges from 17.63 cm to 48.60 cm. Peduncle length ranges from 18.00 cm to 78.50 cm and the diameter of the third internodes from 0.55 cm to 1.30 cm. As for the leaf length and leaf width of the third internodes they vary from 41.85 cm to 98.89 cm and from 3.55 cm to 8.83 cm respectively while the panicle exsertion ranges from 0.00 cm to 34.37 cm. The number of leaves per plant varies from 16 cm to 28 cm while the number of effective tillers was 1 or 2. The weight of 1000 grains accessions ranges from 7.50 g to 52.80 g. The variables with significant variation were panicle length, peduncle length, leaf length of the third internodes, panicle exsertion and weight of 1000 grains (Fig. 2). Table 3. Performance of genotypes of sorghum accessions (Sorghum bicolor (L.) Moench) analyzed. Quantitative traits Mini- mum Maxi- mum Average Standard deviation Hpl (cm) 213.33 486.36 339.57 60.83 Lpa (cm) 17.63 48.60 33.43 6.83 Lpe (cm) 18.00 78.50 43.89 11.17 Dte (cm) 0.55 1.30 0.93 0.16 Ltf (cm) 41.85 98.89 74.82 9.91 ltf (cm) 3.55 8.83 5.59 1.05 Exp (cm) 0.00 34.37 10.08 6.59 A C B
  • 8. Int. J. Agri. & Agri. R. Missihoun et al. Page 30 Quantitative traits Mini- mum Maxi- mum Average Standard deviation Nfe (Number) 16.00 27.67 20.47 2.35 Nte (Number) 1.00 1.75 1.08 0.17 p~1000 (g) 7.50 52.80 32.63 8.07 Hpl = plant height, Lpa = panicle length, Lpe = peduncle length, Dte = diameter of the third internodes, Ltf = leaf length of the third internodes, ltf = leaf width of the third internodes, Exp = panicle exsertion, Nfe = number of leaves, Nw = number of effective tillers and p = ~ 1000g weight of 1,000 grains. Fig. 2. Global Change in quantitative variables within accessions. Correlation between quantitative variables The correlation matrix between the measured variables is presented in Table 4. The analysis of this table shows that plant height was significantly and positively correlated with other variables except for panicle exsertion and number of effective tillers. The panicle length was positively correlated with the length of the peduncle, the leaf length and leaf width of the third internodes and the weight of 1000 grain (p ~ 1000). Peduncle length was negatively correlated to the diameter of the third internodes and number of leaves but was significantly and positively correlated with panicle exsertion. The diameter of the third internodes was positively correlated with the leaf length and leaf width of the third internodes and the number of leaves but negatively correlated to the panicle exsertion. Leaf length of the third internodes was positively correlated with leaf width, number of leaves and weight of 1000 grains (p ~ 1000) and negatively correlated with panicle exsertion. Leaf width of the third internodes was positively correlated only to the number of leaves. With regard to the panicle exsertion and the effective number of tillers, they were negatively correlated to the number of leaves and the weight of 1000 grains (p ~ 1000) respectively and the positively correlated to the weight of 1000 grains (p ~ 1000). Table 4. Correlation matrix of quantitative variables measured. Hpl Lpa Lpe Dte Ltf ltf Exp Nfe Nte p~1000 Hpl 1.00 Lpa 0.73* 1.00 Lpe 0.31* 0.28* 1.00 Dte 0.41* 0.20 -0.28* 1.00 Ltf 0.53* 0.45* -0.18 0.50* 1.00 ltf 0.43* 0.23* -0.11 0.73* 0.48* 1.00 Exp -0.02 -0.15 0.74* -0.24* -0.39* -0.01 1.00 Nfe 0.28* 0.09 -0.37* 0.44* 0.39* 0.47* -0.27* 1.00 Nte -0.12 -0.14 -0.07 -0.02 0.01 -0.02 -0.02 0.07 1.00 p~1000 0.34* 0.25* -0.11 0.13 0.43* -0.01 -0.32* 0.10 0.24* 1.00 * Significant correlation at 5%. PCA analysis revealed variations within sorghum landraces The principal component analysis (PCA) was used to explain the quantitative variations observed within the plant material used. The correlations between variables and the synthetic axes are showed in Fig. 3. Axes 1 and 2 put together account for 54.91% of the total variability in the sorghum accessions studied (Fig. 3). As for the factorial scheme (axis 1 and axis 3), it stands for 48.50% of the total variability. Overall, the first three axes account for nearly 70% of the total
  • 9. Int. J. Agri. & Agri. R. Missihoun et al. Page 31 variation. These three axes were used to describe the variability within the collection. Fig. 3. Circle of correlation (ACP) in the plane 1-2. Plant height, panicle length, diameter of the third internodes, length and width of the third leaf and the number of leaves were significantly and positively correlated with axis 1. On the other hand, axis 2 was significantly and negatively correlated with the length of the peduncle and panicle exsertion. As for the weight of 1000 grains, this variable was significantly and negatively correlated to the main axis 3 (Table 5). These variables were analyzed and interpreted respectively in accordance with their respective axis. Table 5. Coordinates of variable on the first three axes. Variables axis 1 axis 2 axis 3 Hpl 0.700796* -0.595646 -0.122998 Lpa 0.559952 -0.562835 -0.312229 Lpe -0.278482 -0.904343* -0.017289 Dte 0.762000* 0.091818 0.397449 Ltf 0.826384* -0.024270 -0.168076 Ltf 0.702494* -0.091277 0.555841 Exp -0.464255 -0.642451 0.366395 Nfe 0.620137 0.283847 0.295721 Nte -0.003610 0.241112 -0.281302 p~1000 0.437383 0.049252 -0.714529* Grouping accessions on the basis of quantitative and qualitative variables A hierarchical classification of sorghum accessions studied and performed with the aggregation criteria of Ward from adjusted mean values for 15 variables (10 quantitative and qualitative 5) from analysis of variance results in the dendrogram (Fig. 4). The Analysis of this dendrogram was able to differentiate four main groups by truncation at a distance of aggregation of 19. Group I contains 13 accessions that were mainly from durra race with bitter or no bitter yellow grains (SOD 40, SOD 41, SOD 42, SOD 44 and SOD 60), an accession with yellowish grains of caudatum race (SOD 71) and an accession of coloring sorghum of intermediate race durra-kafir (SOD 6). Group II composes of 24 accessions which were mainly sorghum accessions of guinea race with long and white grains of which vegetative cycle was long (SOD 21, SOD 22, SOD 33, SOD35, SOD 37, SOD 47, SOD 57 and SOD 62) and also contains some accessions with red grains as SOD 2, SOD 4 and SOD 65 (all belonging to guinea race). Group III consists of 20 accessions which were essentially guinea race with white or whitish grains and with medium growth cycle (SOD 24, SOD 29, SOD31, SOD 36, SOD54, SOD 56, SOD59, and SOD 61). The accession of the bicolor race (SOD 48) and an accession of caudatum race (SOD 66) were also found in this group. Finally, group VI consists of 17 accessions, mainly early guinea race with red grains of small size (SOD 7, SOD 10, SOD 13, SOD 15, SOD 16, and SOD 64) but also some accessions with white grains (SOD 17, SOD 26 and SOD 38). Fig. 4. Ascending hierarchical classification (AHC) of 74 sorghum accessions according to the Ward aggregation criteria.
  • 10. Int. J. Agri. & Agri. R. Missihoun et al. Page 32 Racial classification of sorghum accessions 76 sorghum accessions collected belong to four botanical races (guinea, Durra, caudatum and bicolor) out of the existing five races. The guinea race was predominant and represents 86.84% of the collection followed by durra race with 5.26%, caudatum race with 2.63% and bicolor race with 1.32%. The intermediaries represent 3.95% (Fig. 5, Table 1). In terms of grain color, 41.33% had an intermediate color (pink, reddish, whitish, yellowish, blackish, white-spotted purple or red, multicolored, etc.), 30.66% of the grains were white, 21.33% were red and 6.66% were yellow. Moreover, it should be noted that 53.95% of accessions had black glumes, 80.26% had loose panicles and 42.11% had a short vegetative cycle. Fig. 5. Proportional distribution of different botanical races of accessions analyzed. Discussion Multivariate analysis of quantitative and qualitative data revealed a structuring in 4 distinct groups of sorghum accessions based mainly on racial factor, but also the duration of the vegetative cycle and morpho- physiological characteristics of grains (color, size, degree of bitterness). This structuring of Benin accessions according to botanical race confirms once again previous agro-morphological investigations on collections from countries of West Africa (Deu et al., 2006; Barro-Kondombo et al., 2008.). The structuring according to morpho-physiological characteristics of the grain was also reported in some studies such as those conducted by Barro-Kondombo et al. (2008) who identified, after agro-morphological characterization, an original group of accessions of local varieties of sorghum with red grains. In the present study, the group of accessions that appears more homogeneous on the basis of grain color is group I which consists of accessions with bitter or no bitter yellow grains. This phenotypic difference of this group of accessions appears extremely important for the judicious use in research because sorghum local varieties with bitter yellow grains are widely recommended for use in traditional medicine in Benin (Kayodé et al., 2005; Missihoun et al., 2012a). In addition, groups II, III, and IV compose of landraces of guinea race which differ in the duration of vegetative cycle and the nature of the grains. These two factors also seem to play an important role in agro-morphological structuring of accessions of cultivated sorghum in Burkina Faso (Barro- Kondombo et al., 2008). However, in Benin, ethno- botanical investigations revealed the specific importance of each of these different groups of local varieties. Local varieties with red grains and short growth cycle (group IV) currently dominate Benin sorghum due to their relative tolerance to drought, their earliness (early maturity), their adaptability to relatively poor soils and their high market value. Local varieties with white grains or intermediate color, medium or long growth cycle (group II and group III) are often more profitable in terms of production of grains, more farinaceous and are used in the production of local dough (Wô, Dibou, Sifanou, Foura). They are sensitive to drought and require soil fertility. Finally, local varieties bitter yellow grains (group I) were mainly grown for use in traditional medicine. This structuring observed with the agro- morphological traits was globally confirmed by molecular analyzes with nuclear microsatellite markers used on the same collection (Missihoun et al., 2015). In terms of selection and/or varietal improvement, positive correlations observed between the different quantitative variables showed that the accessions with long panicles were of big size (big plant height) and those with the weight of 1000 grains had the largest
  • 11. Int. J. Agri. & Agri. R. Missihoun et al. Page 33 diameter of the third internodes. On-farm, farmers had a preference for varieties of high size due to both the quality of their food grains and especially to their important role they play in the production of biomass for combustion and building materials (Kayodé et al., 2006). The utilization of these results will be very important in plant breeding programs to improve the performance of local varieties in terms of grain and biomass production. The racial diversity of cultivated sorghum in Donga department is very high (presence of 4 races out of the five existing races) with a predominance of guinea race. The dominant presence of guinea race in the regions of West Africa is known recognized and has been reported by many researchers from West African countries such as Burkina Faso (Ollitrault 1987; Zongo, 1991; Barro-Kondombo et al., 2010), Mali (Sagnard et al., 2008), Ghana (Kudadjie, 2006) and even in Central Africa with Cameroon (Barnaud, 2007). The present study is consistence with those reported above. The results of this work in Benin could be justified in terms of the geographical position of the country in relation to the history of diversification of sorghum. As a matter of fact, Benin is located in the central of West African countries where there is the diversification of guinea race and with high likelihood of plant materials (seeds) exchanges between farmers of the neighboring countries such as Burkina Faso and Niger. The predominance of guinea race could also be explained by climatic factor. Guinea race would be more suited to the higher rainfall areas (Barro-Kondombo et al., 2010). Furthermore, the absence of kafir race among collected samples in Benin confirms all the previous work done in the regions of West Africa (Burkina Faso, Niger, and Ghana). Deu et al. (2008) observed the same racial diversity in Niger (all races except Kafir) with the difference that their proportion were not the same: durra (23.1%), caudatum (21.3%), guinea (10.5%) and bicolor (8.1%). Kafir race is reported to be found in southern Africa and is almost absent in the regions of West Africa, although more intermediate of this race with other races exist. Conclusion This study is one of the first evaluations which focused on agro-morphological characteristics of grown sorghum showed an important phenotypic variability of local varieties of sorghum kept and managed by farmers in Donga. Based on the traits studied four groups structured around factors such as morpho-physiological characteristics of the grains (color, size, taste) and race, were identified. Moreover, the results showed the presence of four pure races (guinea, Durra, caudatum bicolor) and the intermediaries such as guinea-durra, guinea-kafir, kafir-durra in the surveyed areas. Guinea race is predominant among the races. The use of molecular genetic markers in further work will allow thorough analysis of the evolution factors of genetic diversity for a better use of local resources of grown sorghum in breeding and conservation programs. The interest in the long run, is the sustainability of sorghum that will continue to play its role in food security and the diversification of cereal production country. Acknowledgment The authors thank Mrs Avohou Germaine for her help for collecting data. We thank also the anonymous reviewers for their valuable comments. Adoukonou-Sagbadja H, Dansi A, Vodouhè R, Akpagana K. 2006. Indigenous knowledge and traditional conservation of Fonio millet (Digitaria exilis Stapf, Digitaria iburua Stapf) in Togo. Biodiversity and Conservation 15, 2379-2395. Barnaud A, Deu M, Garine E, McKey D, Joly HI. 2007. Local genetic diversity of sorghum in a village in northern Cameroon: structure and dynamics of landraces. Theoretical and Applied Genetics 114, 237- 248. Barro-Kondombo C, Sagnard F, Chantereau J, Deu M, vom Brocke K, Durand P, Gozé E, Zongo JD. 2010. Genetic structure among sorghum landraces as revealed by morphological variation and microsatellite markers in three agroclimatic regions
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