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African Crop Science Journal
African Crop Science Society
ISSN: 1021-9730 EISSN: 2072-6589
Vol. 8, Num. 2, 2000, pp. 137-143
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African Crop Science Journal, Vol. 8. No. 2, pp. 137-143
African Crop Science Journal, Vol. 8. No. 2, pp. 137-143
PHENOTYPIC DIVERSITY IN THE ETHIOPIAN NOUG GERMPLASM
Tsige Genet and Ketema Belete1 Adet Research Centre, P.O. Box 8, Dahir Dar, Ethiopia 1Alemaya University of Agriculture, P.O. Box, 138, Dire Dawa, Ethiopia
(Received 31 March, 1999; accepted 27 February, 2000)
Code Number: CS00014
INTRODUCTION
Noug (Guizotia abyssinica (L.f.) Cass; 2n = 30) belongs
to the family Compositae and the genus Guizotia which is small with only
six species, of which five are native to Ethiopia (Baagoe, 1974). Guizotia
abyssinica is believed to have originated through disruptive selection from
Guizotia scabra, a common weed in Ethiopia (Doggett, 1987). Noug is extensively
cultivated in Ethiopia and India. It is also grown in Sudan, Uganda, Zaire,
Tanzania, Malawi, Zimbabwe, the West Indies, Nepal, Bangladesh, and Bhutan (Weiss,
1983).
Noug contributes towards 50 - 60% of the Ethiopias edible
oil needs (Riley and Belayneh, 1989). Noug is grown between 1500 - 2400 m above
sea level, in rotation with cereals on poorly drained soils. Gojjam, Shoa, Gonder
and Welega are the largest areas of production, accounting for about 90% of
the countrys total production (Getinet and Alemayehu, 1997). Noug has a reputation
for its ability to grow on marginal soils and to withstand waterlogging because
of its tolerance to low oxygen level (Mesfin, 1975). Fertiliser and drainage
promotes luxurious vegetative growth but do not increase seed yield (Riley and
Belayneh, 1989). Different types of noug are known to be widely cultivated in
Ethiopia (Getinet and Sharma, 1996). The predominant type is 'abat' noug grown
during the main season from June to December. This type has higher yields, and
a high oil content. 'Bungne' noug is adapted to lowland and matures earlier
than 'abat', but is lower yielding and bears lighter seed with lower oil content.
A third type of noug, 'mesno' is tolerant to frost and grows on residual moisture.
Noug seed yields a pale yellow oil with a pleasant nutty taste.
The seed contains about 40% oil with a fatty acid composition of 75 - 80% linoleic
acid (Getinet and Tekelewold, 1995), which makes it important in preventing
arteriosclerosis in humans (Vaughan, 1970). Noug oil is also used for cooking,
as an illuminant, as a lubricant and for manufacturing soap and paints (Simmonds,
1976). It also has potential for cosmetics (Vaughan, 1970).
Noug germplasm has been collected from farmers fields and
is mostly conserved and documented at the Biodiversity Institute of Ethiopia
(the then Plant Genetic Resources Centre for Ethiopia). The distribution and
degree of phenotypic diversity of noug in different agro-ecological areas of
the country has, however, not yet been studied. Hence, this study was undertaken
to estimate the phenotypic diversity in the Ethiopian noug germplasm collections
across a wide range of characters.
MATERIALS AND METHODS
The study was conducted at Adet Research Centre (longitude 37° 29E and latitude 11°16N and 2240 metres above sea level (masl)) located in north western Ethiopia during the 1993 main cropping season (Keremet). For this study, 157 noug accessions were obtained from the Biodiversity Institute of Ethiopia. These were randomly sampled from varying agro-ecological areas of the country. Forty seeds per accession were planted on vertisols in a single row of 4 m length plot, with a spacing of 1 m between rows, and 10 cm between plants. Fertiliser (both N and P2O5) was applied at the rate of 23 kg ha-1 each at the time of planting. Weeding was carried out 21 days after emergence and 45 days after the first weeding.
Observations on quantitative (days to flowering, days to maturity,
and plant height) and qualitative characters (leaf colour, leaf margin, stem
colour, stem hairiness, and angle of branching) were recorded following the
noug description format of the Biodiversity Institute of Ethiopia (Table 1).
Days to 50% flowering is the actual count of the number of days from planting
to the date at which about 50% of the plants in a plot have the first flower.
Days to maturity is the actual count of the number of days from planting to
the date at which 95% of the plants in a plot have reached physiological maturity.
Plant height is the actual measurement in cm taken from the ground to upper
most leaf at maturity. Leaf colour is the most predominant colour of the leaf
during 50% flowering. Allowable states are light green, green, and dark green.
Leaf margin is a visual measurement taken at the middle part of the plant at
50% flowering. Allowable states are smooth, pointed, and round. Stem colour
is the most predominant colour of the stem recorded at 50% flowering. Allowable
states are green and purple. Stem hairiness is a visual measurement at the middle
part of the stem during 50% flowering. Allowable states are hairy, intermediate,
and very hairy. Angle of branching is a visual measurement taken at 50% flowering
at the middle part of the plant. Allowable states are acute and hanged. Days
to 50% flowering, days to maturity, and plant height with continuous variation
were included in order to examine their value in diversity studies.
Table 1. List of characters and their respective classes used
for the evaluation of phenotypic diversity of noug germplasm at Adet, Ethiopia
in 1993 |
Character |
Abbreviations |
Character states |
Codes used |
Phenological |
Days 50% flowering |
DF |
< 75 days
76 - 90 days
91 - 105 days
106 - 120
> 121 days |
1
2
3
4
5 |
Days to maturity |
DM |
< 100 days
101 - 115 days
116 - 130 days
131 - 145 days
> 146 days |
1
2
3
4
5 |
Morphological
|
Leaf colour
|
LC |
Light green
Green
Dark green |
1
2
3 |
Leaf margin
|
LM |
Smooth
Round
Pointed |
1
2
3 |
Stem hairiness |
SH |
Less hairy
Intermediate
Very hairy |
1
2
3 |
Stem colour |
SC |
Green
Purple |
1
2 |
Angle of branching
|
AB |
Acute
Hanged |
1
2 |
Plant height |
PH |
< 100 cm
101 - 130 cm
131 - 160 cm
161 - 190 cm
> 191 |
1
2
3
4
5 |
The quantitative characters which are scaled randomly are useful
in this study, as the noug accessions show considerable variation within and
between regions for these characters. Quantitative characters are in general
of more interest to the plant breeder than the discrete or qualitative characters.
The phenotypic frequencies of the characters were analysed by the Shannon-Weaver
diversity index (H) as described by Jain et al. (1975) in order to estimate
the diversity of each character within each province. For this purpose, the
country was divided into four regions: northern Ethiopia (Tigrai, Gonder, and
Welo), western Ethiopia (Gojjam, Welega, and Ilubabor), southern Ethiopia (Gamu
Gofa, Sidamo, and Bale), and central and eastern Ethiopia (Shewa, Arsi, and
Hararghe).
The Shannon-Weaver diversity index described by Jain et
al. (1975) was calculated as:
H=-SPilog2Pi
Where, Pi is the proportion of the accessions in the ith class of an n class trait in a population. This index has been extensively used to estimate phenotypic diversity in germplasm collections (e.g. Jain et al.,
1975; Tolbert et al., 1979). The additivity of the H allows characters
to be pooled over provinces and over regions (Tolbert et al., 1979).
This property permits the use of hierarchical analysis of variance for testing
the significance of various components of variation in H (regions, provinces,
and characters). Each H value was standardised by dividing it by its maximum
value (log2 n), which ensured that all scaled H values were in the range of 0 to 1.
The minimum value of the index is zero for a monomorphic population.
The value of the index increases with the increase in polymorphism and reaches
the maximum value when all phenotypic classes have equal frequencies (Yang et
al., 1991).
The variance of H has not been characterised. However, assuming
that the eight characters used in our study represent a random sample of all
possible characters of noug plant, an empirical variance was computed from the
eight estimates of Shannon-Weavers diversity index.
RESULTS AND DISCUSSION
Distribution of phenotypic classes. The phenotypic frequencies
for individual characters and provinces as percentage of the number of accessions
from each geographic origin are presented in Table 2. The area of origin were
grouped into regions and regions into provinces and weighted regional frequencies
were computed.
Table 2. Phenotypic diversity of noug based on percentage of
accessions for each province |
Geographic origin |
Max-N*
N |
Leaf colour |
Leaf margin |
Stem hairiness |
Stem colour |
Angle of branching |
Days to flowering |
Days to maturity |
Plant height |
1 |
2 |
3 |
1 |
2 |
3 |
1 |
2 |
3 |
1 |
2 |
1 |
2 |
1 |
2 |
3 |
4 |
5 |
1 |
2 |
3 |
4 |
5 |
1 |
2 |
3 |
4 |
5 |
Northern Ethiopia |
Tigrai |
9 |
22 |
67 |
11 |
11 |
0 |
89 |
0 |
89 |
11 |
0 |
100 |
100 |
0 |
22 |
67 |
0 |
11 |
0 |
0 |
67 |
33 |
0 |
0 |
0 |
78 |
22 |
0 |
0 |
Gonder |
41 |
78 |
15 |
7 |
0 |
2 |
98 |
80 |
20 |
0 |
2 |
98 |
80 |
20 |
0 |
62 |
34 |
4 |
0 |
2 |
44 |
37 |
12 |
5 |
0 |
2 |
54 |
44 |
0 |
Welo |
13 |
69 |
31 |
0 |
0 |
0 |
100 |
31 |
69 |
0 |
0 |
100 |
54 |
46 |
23 |
69 |
8 |
0 |
0 |
8 |
54 |
15 |
23 |
0 |
2 |
20 |
51 |
27 |
0 |
Region |
63 |
68 |
25 |
7 |
2 |
2 |
96 |
59 |
40 |
1 |
2 |
98 |
78 |
22 |
8 |
65 |
25 |
2 |
0 |
3 |
49 |
32 |
13 |
3 |
2 |
20 |
41 |
27 |
0 |
Western Ethiopia
|
Gojam |
28 |
71 |
29 |
0 |
4 |
0 |
96 |
46 |
54 |
0 |
4 |
96 |
75 |
25 |
4 |
53 |
32 |
7 |
4 |
8 |
25 |
46 |
21 |
0 |
0 |
10 |
61 |
29 |
0 |
Welega |
26 |
77 |
19 |
4 |
4 |
4 |
92 |
62 |
38 |
0 |
58 |
42 |
88 |
12 |
0 |
69 |
27 |
4 |
0 |
0 |
46 |
38 |
16 |
0 |
0 |
8 |
65 |
27 |
0 |
Ilubabor |
3 |
67 |
33 |
0 |
0 |
0 |
100 |
100 |
0 |
0 |
0 |
100 |
100 |
0 |
0 |
67 |
0 |
33 |
0 |
0 |
67 |
33 |
0 |
0 |
0 |
33 |
67 |
0 |
0 |
Region |
57 |
74 |
25 |
1 |
4 |
2 |
94 |
56 |
44 |
0 |
30 |
70 |
82 |
18 |
2 |
65 |
28 |
3 |
2 |
4 |
37 |
42 |
17 |
0 |
0 |
11 |
63 |
26 |
0 |
Southern Ethiopia
|
Gamu Gofa |
2 |
100 |
0 |
0 |
0 |
0 |
100 |
50 |
50 |
0 |
0 |
100 |
0 |
100 |
0 |
0 |
50 |
50 |
0 |
0 |
0 |
50 |
50 |
0 |
0 |
0 |
50 |
50 |
0 |
Sidamo |
1 |
0 |
100 |
0 |
0 |
0 |
100 |
100 |
0 |
0 |
0 |
100 |
100 |
0 |
0 |
0 |
100 |
0 |
0 |
0 |
100 |
0 |
0 |
0 |
0 |
0 |
100 |
0 |
0 |
Bale |
4 |
0 |
100 |
0 |
0 |
0 |
100 |
75 |
25 |
0 |
2 |
98 |
75 |
25 |
0 |
100 |
0 |
0 |
0 |
0 |
75 |
25 |
0 |
0 |
0 |
25 |
75 |
0 |
0 |
Region |
7 |
57 |
43 |
0 |
0 |
0 |
100 |
71 |
29 |
0 |
14 |
85 |
57 |
43 |
0 |
57 |
29 |
14 |
0 |
0 |
71 |
29 |
0 |
0 |
0 |
14 |
57 |
29 |
0 |
Central and E. Ethiopia
|
Shewa |
23 |
70 |
26 |
4 |
0 |
0 |
100 |
43 |
48 |
9 |
4 |
96 |
78 |
22 |
17 |
78 |
5 |
0 |
0 |
4 |
48 |
31 |
17 |
0 |
4 |
39 |
52 |
5 |
0 |
Arsi |
6 |
83 |
17 |
0 |
0 |
0 |
100 |
83 |
17 |
0 |
0 |
100 |
83 |
17 |
0 |
100 |
0 |
0 |
0 |
6 |
61 |
33 |
0 |
0 |
0 |
17 |
66 |
17 |
0 |
Shararghe |
1 |
0 |
100 |
0 |
0 |
0 |
100 |
0 |
100 |
0 |
0 |
100 |
100 |
0 |
0 |
0 |
0 |
100 |
0 |
0 |
100 |
0 |
0 |
0 |
0 |
0 |
100 |
0 |
0 |
Region |
30 |
70 |
27 |
3 |
0 |
0 |
100 |
50 |
43 |
7 |
3 |
97 |
80 |
20 |
13 |
80 |
3 |
4 |
0 |
3 |
53 |
30 |
14 |
0 |
3 |
33 |
56 |
8 |
0 |
Ethiopia |
157 |
70 |
26 |
4 |
2 |
1 |
97 |
56 |
42 |
2 |
13 |
87 |
79 |
21 |
6 |
66 |
22 |
5 |
1 |
3 |
47 |
35 |
14 |
1 |
1 |
18 |
57 |
24 |
0 |
Max - N = Maximum number of accessions in each province;
See Table 1 for the description of numbers which are below each characters
|
Provinces representing small number sampled showed high monomorphism,
namely Gamu Gofa, Sidamo and Hararghe. Thus, variability for the eight characters
examined were widely distributed. Individual characters differed in their patterns
of distribution and amount of variation.
Leaf colour is widely distributed throughout the collection,
i.e., 70% light green, 26% green, and 4% dark green. Except for Gamu Gofa, Sidamo,
Bale, and Hararghe, the rest of the provinces have intermediate frequencies
in leaf colour. The collection had predominantly pointed type of leaf margin
in all provinces.
The accessions were dominated by less hairy stem (56%) and
intermediate proportions of hairy stem (42%). Very hairy phenotypes were detected
in Tigrai and Shewa provinces only. Gojjam had almost equal proportion of less
hairy and intermediate characters, while very hairy phenotypes were not detected.
Although environmentally sensitive, purple stem colour was
the dominant colour noted in the accessions. Welega province had the highest
frequency of green stem colour. The two phenotypic classes of angle of branching
were observed at intermediate frequencies only in Welo province (Table 2). Acute
type of branching was a more common phenotype. The continuous metric traits,
such as days to flowering, days to maturity and plant height, had intermediate
frequencies in their measurement.
Analysis of phenotypic diversity index (H). Table 3
gives the estimate of H individually and pooled over characters and provinces
and appropriately weighted by number of accessions. The three highest mean values
of H were from Gojjam (0.60), Welega (0.57), and Shewa (0.53) provinces. The
lowest values of the H were from Sidamo and Hararghe (0.00), Gamu Gofa (0.24),
and Ilubabor (0.23). One possible reason for these low H values could have
been the small number of samples for the study from these provinces. On the
other hand, there might be greater natural selection for noug in both provinces
due to generally low rainfall in the north and high rainfall in the south. Clearly,
these results indicate that the amount of variation differ considerably among
the provinces.
Table 3. Estimates of phenotypic diversity for 8 characters
used for the evaluation of noug germplasm at Adet, Ethiopia, in 1993 |
Gepgraphic origin |
Morphological and phenological characters* |
LC |
LM |
SH |
SC |
AB |
DF |
DM |
PH |
H ± S3 |
Northern Ethiopia |
Tigrai |
0.77 |
0.32 |
0.32 |
0.00 |
0.00 |
0.52 |
0.39 |
0.00 |
0.33 |
0.08 |
Gonder |
0.60 |
0.09 |
0.46 |
0.14 |
0.72 |
0.44 |
0.75 |
0.48 |
0.46 |
0.08 |
Welo |
0.57 |
0.00 |
0.57 |
0.00 |
1.00 |
0.49 |
0.72 |
0.68 |
0.50 |
0.12 |
Region |
0.72 |
0.18 |
0.66 |
0.14 |
0.76 |
0.56 |
0.74 |
0.68 |
0.56 |
0.08 |
Western Ethiopia
|
Gojjam |
0.55 |
0.16 |
1.00 |
0.24 |
0.81 |
0.71 |
0.77 |
0.55 |
0.60 |
0.09 |
Welega |
0.53 |
0.30 |
0.60 |
0.98 |
0.53 |
0.46 |
0.63 |
0.52 |
0.57 |
0.06 |
Ilubabor |
0.58 |
0.00 |
0.00 |
0.00 |
0.00 |
0.39 |
0.39 |
0.39 |
0.23 |
0.08 |
Region |
0.56 |
0.24 |
0.62 |
0.91 |
0.68 |
0.56 |
0.72 |
0.55 |
0.61 |
0.06 |
Southern Ethiopia
|
Gamu Gofa |
0.00 |
0.00 |
0.63 |
0.00 |
0.00 |
0.43 |
0.43 |
0.43 |
0.24 |
0.08 |
Sidamo |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
Bale |
0.00 |
0.00 |
0.51 |
0.14 |
0.81 |
0.00 |
0.35 |
0.35 |
0.27 |
0.09 |
Region |
0.62 |
0.00 |
0.55 |
0.58 |
0.99 |
0.59 |
0.37 |
0.59 |
0.54 |
0.09 |
Central and Ethiopia
|
Shewa |
0.66 |
0.00 |
0.85 |
0.24 |
0.76 |
0.40 |
0.71 |
0.61 |
0.53 |
0.09 |
Arsi |
0.42 |
0.00 |
0.42 |
0.00 |
0.66 |
0.00 |
0.52 |
0.54 |
0.32 |
0.09 |
Hararghe |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
0.00 |
Region |
0.64 |
0.00 |
0.82 |
0.19 |
0.72 |
0.42 |
0.67 |
0.62 |
0.51 |
0.09 |
Ethiopia |
0.45 |
0.08 |
0.50 |
0.22 |
0.53 |
0.37 |
0.51 |
0.46 |
0.40 |
0.05 |
* LC = Leaf colour, LM = Leaf margin, SH = Stem hairiness,
SC = Stem colour, AB = Angle of branching, DF = Days to 50% flowering,
DM = Days to maturity, PH = Plant height, H = Diversity index
|
The estimate of diversity index for individual characters pooled
over provinces and regions showed considerable variation among characters (Table
3). For example, the index varied from 0.98 for leaf margin to 0.53 for angle
of branching country-wide. The average diversity across 8 characters from all
the regions or the overall diversity for Ethiopia was H = 0.40. Based on the
average diversity calculated from the individual regions across all characters,
the most diverse region was western Ethiopia H = 0.61 and the least diverse
was central and eastern Ethiopia H 0.51. Generally, northern Ethiopia is considered
to be the centre of origin and diversity of noug (Baagoe, 1974), but in this
analysis northern Ethiopia had estimates of H = 0.56. This might be due to
natural selection in the areas of northern Ethiopia which are frequently affected
by drought and/or it could also be due to sampling effect.
Hierarchical analysis of variance in H values for individual
characters (Table 4) shows that diversity among regions was not significantly
different (P>0.05), whereas greater variance was obtained among provinces
within different characters (P<0.05).
Table 4. Hierarchical analysis of variance for
diversity index (H) |
Source |
DF |
SS |
MS |
F |
P% |
Expected mean square |
Between regions |
3 |
1 |
0.44 |
1.52 |
0.2846 |
d2c + cd2b
+ cb S2a/a-1 |
Between provinces within regions |
8 |
2 |
0.29 |
4.74 |
0.0159 |
d2c + cd2b |
Characters within provinces
|
84 |
5 |
0.06 |
|
|
d2c |
Total |
95 |
|
|
|
|
|
S2a
= added variance component due to region
d2b = variance due to provinces within
regions
d2c = variance due to characters within
provinces
a = number of regions
b = number of provinces
c = number of characters
n = (abc) = total number of sample sizes = 4 x 3 x 8 = 96 |
In conclusion, based on limited characters studied and small
collections evaluated, the centres of diversity for noug appears to be in Gojjam,
Welega, and Shewa. Further exploration should be undertaken in these regions.
The utility of germplasm collection to research programmes
designed to locate genes depends on adequate sampling procedures. The Ethiopian
collection presently has a large number of accessions representing a valuable
reservoir of genetic resources. However, several important areas in Ethiopia
are not still adequately represented. Hence, several noug collection missions
representing diverse agro-ecological areas of the country should be organised
for carrying out diversity analysis to derive some guidelines for conservation
activities.
ACKNOWLEDGEMENTS
This work was supported by the International Development Research
Centre (IDRC) of Canada. The authors acknowledge the help of Dr. Getinet Alemaw
and Dr. P.A. Chadhokar in reviewing the manuscript, and thank Alamine Atanaw,
Melkie Nibret, and Menychel Alameneh for their assistance in the fieldwork.
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