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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
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 Ethiopia’s 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 country’s 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° 29’E and latitude 11°16’N 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-Weaver’s 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.

REFERENCES

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