Introduction
Vicia L. is a genus with around 232 species in the world and 45 species in Iran, from the legume family, Fabaceae, as an annual and perennial herb. These species have been known by the common name vetches. The genus is primarily found in the Mediterranean and Irano-Turanian regions, such as in Iran, Anatoly, Caucasus, Iraq, Afghanistan, Central Asia, Talesh, Syria, Armenia, Turkmenistan, Jordan, North Africa, Greece, Pakistan, and Palestine1. Vetches are short-lived forage plants that are highly resistant to cold and dehydration conditions and can be grown in rainfed and irrigated climates. They fix nitrogen in the soil by fixation in root nodes, and help to soil erosion by planting in sloping areas2,3. As a legume crop, it provides nitrogen to the soil and reduces the incidence of diseases in succeeding non-leguminous crops. Their widespread adaptation and excellent capacities to produce biomass make them very attractive to farmers4. One attraction of vetch is its versatility, which permits diverse utilization as either ruminant feed or green manure. Because of rapid growth in the first year, different species of Vicia spp. can be used to improve overall livestock, feed quality, improve soils, agriculture for fodder, green manure, human nutrition, and the pharmaceutical industry5.
Iran is a genetics resources of the genus Vicia and it is widely distributed in different habitats and conditions. Most of the plants in Vicia genus show more variety in morphological traits and sometimes it is difficult to distinguish species of this genus6,7.
Genetic variation among Vicia genotypes is imperative for their efficient utilization in plant breeding schemes and effective conservation. Diversity studies available in germplasm, collections have been performed on many plant species for Vicia genus from different regions of the world. In comparison to other annual forage legumes, advances in breeding vetches (Vicia spp.) are rather modest. It has been one of the morphological characteristics of the plant reported in V. sativa8-14, V. faba15, V. narbonensis8,10,11,15, V. ervilia16, V. villosa10,11, V. atropurpurea11, V. dasycarpa8, V. hybrid, V. pannonica, V. lutea, V. peregrine, V. lathyroides and V. grandiflora11.
There are 335 accessions of 25 Vicia spp. in natural resources gene bank of Iran, that have been collected from different geographical regions of Iran. In this study, it was aimed to determine some morphological characteristics and forage yields of different vetch genotypes by collecting from natural flora of Iran region. The present study was focused on the estimation of genotypic variation for 12 morphological traits within the V. michauxii, V. michauxii var.stenophylla, V. monantha, V. narbonensis, V. sativa with three varieties: V. sativa var.angustifollia, V. sativa var.cordata, V. sativa var.sativa and V. villosa, to assess their breeding potential and suitability for developing novel common vetch lines with improved agronomic characteristics related to grain production and quality.
Material and methods
Germplasm
A total of 58 germplasm populations were evaluated in this study. This consisted of 1 V. michauxii, 1 V. michauxii var.stenophylla, 1 V. monanta, 1 V. narbonensis, 34 V. sativa, 9 V. sativa var.angustifollia, 1 V. sativa var.cordata, 4 V. sativa var.sativa and 6 V. villosa. The populations were acquired from the Natural Resources Genebank of Iran (Table 1).
Taxon | Code | Abbre. code |
Origin, province | Longitude | Latitude | Altitude (m asl) | ||
---|---|---|---|---|---|---|---|---|
V. michauxii | 2944 | Vmi | East Azerbaijan, Kaleybar | 47° | 02´ | 38° | 51´ | 1500 |
V. michauxii var.stenophylla | 37129 | Vmis | Qom | 50° | 56´ | 34° | 11´ | 2482 |
V. monantha | 32845 | Vmo | Kermanshah | 47° | 14´ | 34° | 8´ | 1338 |
V. narbonensis | 34878 | Vn | Lorestan, Aleshtar | 48° | 10´ | 33° | 45´ | 1495 |
V. sativa | 5321 | Vs | East Azerbaijan | 46° | 16´ | 37° | 54´ | 1750 |
6646 | Vs | Lorestan, Kohdasht | 33° | 40´ | 47° | 30´ | 1200 | |
6654 | Vs | Lorestan, Kohdasht | 33° | 17´ | 47° | 27´ | 1130 | |
6681 | Vs | Lorestan, Kohdasht | 33° | 32´ | 47° | 37´ | 1260 | |
11760 | Vs | Gilan, Rezvanshahr | 37° | 31´ | 49° | 13´ | 280 | |
11761 | Vs | Gilan, Rasht | 36° | 51´ | 49° | 37´ | 80 | |
11762 | Vs | Gilan, Rezvanshahr | 37° | 37´ | 49° | 07´ | 280 | |
11763 | Vs | Gilan, Rasht | 37° | 59´ | 49° | 33´ | 100 | |
11764 | Vs | Gilan, Talesh | 37° | 32´ | 45° | 55´ | 280 | |
11771 | Vs | Gilan, Talesh | 37° | 42´ | 48° | 55´ | 150 | |
11772 | Vs | Gilan, Rezvanshahr | 37° | 32´ | 49° | 07´ | 20 | |
11774 | Vs | Gilan, Rasht | 37° | 11´ | 49° | 39´ | 120 | |
24062 | Vs | Gilan, Astaneh Ashrafiyyeh | 37° | 20´ | 49° | 47´ | 25 | |
24069 | Vs | Gilan, Chabuksar | 36° | 56´ | 50° | 32´ | 170 | |
24074 | Vs | Gilan, Astaneh Ashrafiyyeh | 37° | 19´ | 50° | 07´ | 16 | |
24076 | Vs | Gilan, Chabuksar | 36° | 57´ | 50° | 35´ | 210 | |
24084 | Vs | Gilan, Rahimabad | 37° | 02´ | 50° | 18´ | 40 | |
24097 | Vs | Gilan, Rahimabad | 37° | 01´ | 50° | 17´ | 45 | |
32972 | Vs | Kermanshah, Hersin | 34° | 13´ | 47° | 25´ | 1367 | |
33456 | Vs | Hamadan | 47° | 57´ | 34° | 24´ | 1545 | |
38517 | Vs | Gilan, Siyahkal | 49° | 57´ | 36° | 59´ | 342 | |
38523 | Vs | Gilan, Talesh | 49° | 3´ | 37° | 36´ | 405 | |
38526 | Vs | Gilan | 48° | 46´ | 37° | 41´ | 827 | |
38527 | Vs | Gilan, Astra | 48° | 58´ | 38° | 24´ | 21 | |
38528 | Vs | Gilan, Rudsar | 50° | 12´ | 36° | 48´ | 608 | |
38531 | Vs | Gilan, Rezvan shahr | 49° | 20´ | 37° | 30´ | 315 | |
38532 | Vs | Gilan, Talesh | 49° | 4´ | 37° | 37´ | 450 | |
38533 | Vs* | Gilan | 38° | 10´ | 48° | 20´ | 600 | |
38536 | Vs* | Gilan | 36° | 54´ | 49° | 26´ | 577 | |
40310 | Vs | Kermanshah, Salase babajani | 34° | 49´ | 46° | 05´ | 1395 | |
40315 | Vs | Kermanshah, Salase babajani | 34° | 49´ | 46° | 05´ | 1395 | |
40326 | Vs | Kermanshah, Javanrud | 34° | 48´ | 46° | 33´ | 1525 | |
40334 | Vs | Kermanshah, Salase babajani | 34° | 51´ | 46° | 01´ | 1395 | |
43100 | Vs | Khozestan, Masjed soliman | 31° | 56 | 49° | 18´ | 870 | |
V. sativa var.angustifollia | 38524 | Vsa | Gilan, Siahkal | 50° | 14´ | 36° | 53´ | 670 |
38525 | Vsa | Gilan, Talesh | 48° | 51´ | 37° | 41´ | 281 | |
38530 | Vsa | Gilan, Talesh | 48° | 52´ | 37° | 41´ | 215 | |
38534 | Vsa | Gilan, Rasht | 49° | 35´ | 37° | 0´ | 137 | |
38535 | Vsa | Gilan, Rodbar | 49° | 40´ | 36° | 46´ | 968 | |
38537 | Vsa | Gilan, Gilan | 49° | 31´ | 36° | 56´ | 187 | |
4740 | Vsa | Ilam, Ivan | 46° | 26´ | 33° | 38´ | 1170 | |
7243 | Vsa | Kohkiloye ve Boyerahmad, Firozabad | 52° | 57´ | 28° | 86 | 1900 | |
38529 | Vsa | Gilan, Rezvan shahr | 49° | 5´ | 37° | 28´ | 307 | |
V. sativa var.cordata | 34295 | Vsc | Gilan, Rezvan shahr | 49° | 4´ | 37° | 36´ | 310 |
V. sativa var.sativa | 1862 | Vss | Kermanshah | 47° | 06´ | 34° | 31´ | 1350 |
24631 | Vss | Kermanshah | 47° | 06´ | 34° | 31´ | 1400 | |
29802 | Vss | Kohkiloye ve Boyerahmad | 30° | 59´ | 51° | 07´ | 2380 | |
32900 | Vss | Kermanshah | 34° | 16´ | 46° | 09´ | 1444 | |
V. villosa | 315 | Vv | Alborz, Karaj | 35° | 83´ | 51° | 01´ | 1460 |
322 | Vv | Karaj | 35° | 83´ | 51° | 01´ | 1470 | |
6268 | Vv | Fars, Shiraz, Sepidan, Sheshpir | 30° | 25´ | 51° | 98´ | 2350 | |
14561 | Vv | Merkezi, Arak | 34° | 09´ | 49° | 70´ | 1730 | |
28061 | Vv | Ardabil | 38° | 25´ | 48° | 29´ | 1350 | |
34212 | Vv | Chahar-mahale Bakhtiyari, Borujen | 31° | 46´ | 50° | 59´ | 2600 |
Field trial
Seed of all 58 populations were sown in seedling pots (December 2018). Then planting and maintenance operations were carried out in the field at the research field of Research Institute of Forests and Rangelands, Alborz province, Iran (2018-2020). A week before planting, the soil was prepared as a fine seedbed to enhance good seedling establishment. The field experimental layout was a One-way analysis of variance (ANOVA) designed. The row and plant spacings were 100 and 40 cm, respectively. The trial was managed according to previouse experiences (several hand weeding was practiced, the first hand weeding was made 40 d after crop emergence, and then repeated every forty days until the end of the growing season, to minimize yield reduction due to weed competitions for soil nutrients, water and solar radiation). Irrigation was applied during the trial. The populations were harvested for seed during the period July to November 2020, depending on their maturity.
Morphological traits
Ten plants (normal growth, uniform performance, disease- and insect pest-free) of each 58 Vicia populations were evaluated by 12 different quantitative traits including day to spourat (day to germination), days to first flowering, days to total flowering, days to maturity (days to seed ripening), plant height (at 50 % flowering, cm), internode length (second internode at 50 % flowering, cm), stems number, pod length (cm), pod width (cm), pod index (pod length/width), biomass yield (plant fresh weight) (g), and plant dry weight (g)17.
Data analysis
Data were subjected to analysis of variance (ANOVA) using the SAS software system18. Significant differences among the mean values of 12 traits were compared the DMRT Duncan test. Pearson correlation was determined using SPSS v.21. To evaluate the information contained in the collected morphological data, principal component analysis (PCA) was carried out by Minitab software (version 15). PCA was used to identify the most important traits (plant height, internode length, stems number, pod length, pod width, pod index, biomass yield, dry weight, day to sprout, days to first flowering, days to total flowering, days to maturity) in the data set. Mean values populations were used to create a correlation matrix from which the standardized PCA scores were extracted and a Scatter plot on the first two PCA was performed. Cluster analysis was performed using Ward’s methods and Euclidean distance and a dendrogram was calculated.
Results
The results of analysis variance revealed significant (P<0.01) variation for eight morphological and yield traits among taxa and populations of Vicia spp. except for pod width trait among populations (Table 2). Table 3 shows the comparison of mean morphological and yield traits in nine taxa of Vicia spp. The value of plant height, internode length and stems number differ between 24.50-150 cm, 3.29-15 cm and 2.81-9, respectively. The highest value of plant height (150 cm), stems number (9) and internode length (15 cm), were shown in V. monantha (Vmo) and V. michauxii var. stenophylla (Vmis), respectively. The variation of pod length between taxa was significant and it differs from1.06 cm in V. sativa var.cordata (Vsc) to 4 cm in V. monantha (Vmo). There was no significant difference in pod width between taxa and they located as two groups (a and b), so two taxa of V. michauxii (Vmi and Vmis) had the widest pod (1.14 and 1.1 cm). Despite the significant differences in biomass yield and dry weight traits, V. villosa (Vv) showed the most value of these traits (biomass yield=60.12 g and dry weight=15.63 g).
Source of Variations |
Degrees of freedom (d.f) |
Plant height |
Internode length |
Stem number |
Pod length |
Pod width |
Pod index |
Biomass yield |
Dry weight |
---|---|---|---|---|---|---|---|---|---|
Taxon | 8 | 3770.70** | 48.92** | 25.93** | 5.23** | 0.47** | 9.19** | 3809.63** | 223.70** |
Population | 48 | 905.59** | 12.11** | 15.62** | 1.89** | 0.06 ns | 7.76** | 1967.42** | 202.36** |
Error | 150 | 346.70 | 2.28 | 2.58 | 0.39 | 0.05 | 1.30 | 218.67 | 18.08 |
CV % | 35.88 | 29.18 | 32.38 | 22.77 | 39.08 | 22.53 | 44.56 | 48.83 |
*, ** significant at 0.05 and 0.01 levels, respectively; ns not significant.
Species | Plant height (cm) |
Internode Length (cm) |
Stems number |
Pop length |
Pod width (cm) |
Pod index |
Biomass Yield (g) |
Dry weight (g) |
---|---|---|---|---|---|---|---|---|
V. michauxii (Vmi) | 63.38 cd | 6.56 bc | 2.81 b | 2.54 bc | 1.14 a | 4.36 b | 9.59 cd | 5.46 bc |
V. michauxii var. stenophylla (Vmis) | 100.0 b | 15.0 a | 5.0 b | 2.5 bc | 1.1 a | 2.27 c | 20.0 bd | 5.0 bc |
V. monantha (Vmo) | 150.0 a | 8.0 b | 9.0 a | 4.0 a | 0.6 b | 6.67 a | 0.06 d | 0.01 c |
V.narbonensis (Vn) | 24.50 e | 4.75 cd | 3.0 b | 3.3 ab | 0.65 b | 5.1 ab | 5.13 cd | 1.15 c |
V. sativa (Vs) | 48.41 de | 4.8 cd | 5.08 b | 2.88 bc | 0.56 b | 5.33 ab | 35.06 b | 9.16 ab |
V. sativa var. angustifollia (Vsa) | 45.07 de | 6.9 bc | 4.6 b | 3.27 ab | 0.60 b | 5.48 ab | 19.03 bd | 5.18 bc |
V. sativa var. cordata (Vsc) | 85.13 bc | 8.69 b | 8.75 a | 1.06 d | 0.29 b | 3.75 bc | 27.92 bc | 6.92 bc |
V. sativa var. sativa (Vss) | 54.67 ce | 3.29 d | 4.58 b | 2.0 cd | 0.5 b | 4.0 bc | 41.08 ab | 9.32 ab |
V. villosa (Vv) | 63.56 cd | 3.36 d | 4.5 b | 2.17 c | 0.51 b | 4.26 b | 60.12 a | 15.63 a |
Fifty eight (58) populations of Vicia spp. were compared for vegetative and phenology traits (Table 4). There was a wide range of value in plant height from 19 cm in V. sativa var.angustifollia (38534) to 150 cm in V. monantha (32845), also the most value of plant height between populations of species were shown in V. villosa (322) (100.33 cm). V.michuxii var.stenophylla (37129) (100 cm), V. sativa (38527) (90 cm) and V. sativa var.cordata (34295) (85.13 cm). The length of the internode was a very differet from 1.83 cm in V. sativa (24062, 40334, 43100) to 15 cm in V. michauxii var.stenophylla (37129). Also, 9.83, 8.69, and 8.28 cm of internode length were shown in V. sativa (38527), V. sativa var.cordata (34295), and V. sativa var.angustifollia (38525), respectively. The highest and lowest number of stems were 2 and 15, which were shown in two different taxa of V. sativa species (Vsa38530 and Vs11774). This trait in populations of V. villosa was no significant different. Four populations of V. sativa (38527, 33456, 24074 and 32972), V. sativa var.angustifollia(38525) and V. monantha (32845), had the largest pod in terms of length (4-4.53 cm) and populations V. michauxii var.stenophylla (37129), and V. sativa (5321) had the largest pod in terms of width (1.1 and 1.06 cm). In compare of yield traits (biomass yield and dry weight), three populations of V. sativa: Vs11761, Vs24062, Vs40326, and two populations of V.villosa:Vv322, Vv6268, had the most values of these traits. The values of these traits in these populations were Vs11761 (83 and 26 g.), Vs24062 (83 and 26 g), Vs40326 (103.67 and 36.33 g), Vv322 (108.33 and 38.60) and Vv6268 (83.50 and 19.73 g).
Population | Plant Height (cm) |
Internode Length (cm) |
Stems Number |
Pod Length (cm) |
Pod Width (cm) |
Pod Index |
Biomass Yield (g) |
Dry Weight (g) |
Day to Sprout |
Days to first Flowering |
Days to total Flowering |
Days to Maturity |
|
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Vmi2944 | 78.56 c-e | 7.94 b-e | 2.67 g-i | 2.87 c-j | 0.82 ab | 3.72 k-p | 10.15 j-n | 2.66 i-n | 28 a | 95 a | 115 c | 158 c | |
Vmis37129 | 100 b | 15 a | 5 f-i | 2.5 f-k | 1.1 a | 2.27 op | 20.00 h-n | 5.00 g-n | 28 a | 95 a | 115 c | 158 c | |
Vmo32845 | 150 a | 8 b-e | 9 b-d | 4 a-c | 0.6 bc | 6.67 a-h | 0.06 n | 0.01 n | 28 a | 95 a | 115 c | 158 c | |
Vn34878 | 24.50 i-k | 4.75 f-p | 3.00 g-i | 3.30 b-g | 0.65 bc | 5.10 f-n | 5.13 l-n | 1.16 k-n | 28 a | 95 a | 120 b | 162 b | |
Vs5321 | 43.88 e-k | 4.00 i-p | 2.50 hi | 2.14 g-l | 1.06 a | 2.00 p | 2.07 mn | 0.55 l-n | 28 a | 95 a | 115 c | 158 c | |
Vs6646 | 44.33 e-k | 2.83 n-p | 4.67 f-i | 2.83 c-j | 0.50 bc | 5.67 c-k | 61.83 b-d | 14.61c-f | 28 a | 95 a | 115 c | 158 c | |
Vs6654 | 28.33 h-k | 2.00 p | 4.00 f-i | 2.00 h-l | 0.50 bc | 4.00 j-p | 42.67 e-i | 10.83 d-i | 28 a | 95 a | 115 c | 158 c | |
Vs6681 | 63.00 c-h | 3.33 l-p | 5.00 f-i | 3.97 a-d | 0.50 bc | 7.93 ab | 74.83 bc | 18.43 cd | 28 a | 95 a | 115 c | 158 c | |
Vs11760 | 56.67 c-k | 4.50 g-p | 4.67 f-i | 2.00 h-l | 0.50 bc | 4.00 j-p | 53.00 c-g | 15.27 c-f | 28 a | 95 a | 115 c | 158 c | |
Vs11761 | 66.67 b-g | 6.17 c-l | 5.33 f-h | 2.50 f-k | 0.50 bc | 5.00 f-n | 83.00 ab | 26.00 b | 28 a | 95 a | 115 c | 162 b | |
Vs11762 | 37.67 f-k | 3.67 j-p | 6.00 e-g | 2.50 f-k | 0.50 bc | 5.00 f-n | 19.00 h-n | 5.06 g-n | 28 a | 95 a | 115 c | 158 c | |
Vs11763 | 80.00 b-e | 6.33 c-i | 11 bc | 3.33 b-g | 0.50 bc | 6.67 a-h | 45.00 e-h | 9.33 e-k | 28 a | 95 a | 120 b | 162 b | |
Vs11764 | 61.67 c-i | 5.17 e-o | 11.67 b | 2.67 e-k | 0.50 bc | 5.33 e-m | 51.17 c-g | 9.03 e-l | 28 a | 95 a | 115 c | 158 c | |
Vs11771 | 52.33 d-k | 4.83 f-p | 4.33 f-i | 2.00 f-k | 0.50 bc | 4.00 j-p | 56.67 b-f | 11.43 d-h | 28 a | 95 a | 115 c | 158 c | |
Vs11772 | 65.00 b-h | 3.17 m-p | 4.67 f-i | 2.00 f-k | 0.50 bc | 4.00 j-p | 41.50 e-i | 8.93 e-l | 28 a | 95 a | 120 b | 162 b | |
Vs11774 | 68.33 b-f | 4.33 h-p | 15.00 a | 2.67 e-k | 0.50 bc | 5.33 e-m | 53.00 c-g | 15.27 c-f | 28 a | 95 a | 115 c | 158 c | |
Vs24062 | 40.00 f-k | 1.83 p | 4.67 f-i | 2.00 f-k | 0.50 bc | 4.00 j-p | 83.00 ab | 26.00 b | 28 a | 95 a | 120 b | 162 b | |
Vs24069 | 41.00 f-k | 2.50 n-p | 5.33 f-h | 3.83 a-e | 0.50 bc | 7.67 a-d | 37.17 e-k | 10.43 d-j | 28 a | 95 a | 115 c | 158 c | |
Vs24074 | 49.67 d-k | 4.17 h-p | 6.00 e-g | 4.00 a-c | 0.47 bc | 8.67 a | 44.00 d-h | 10.43 d-j | 28 a | 95 a | 115 c | 158 c | |
Vs24076 | 41.33 f-k | 3.50 k-p | 4.33 f-i | 2.50 f-k | 0.47 bc | 5.50 d-l | 41.50 e-i | 8.93 e-l | 28 a | 95 a | 120 b | 162 b | |
Vs24084 | 55.00 c-k | 5.33 d-o | 4.67 f-i | 2.67 e-k | 0.50 bc | 5.33 e-m | 44.00 d-h | 10.43 d-j | 28 a | 95 a | 115 c | 158 c | |
Vs24097 | 51.33 d-k | 3.00 n-p | 6.00 e-g | 2.00 f-k | 0.50 bc | 4.00 j-p | 45.00 e-h | 8.77 e-m | 28 a | 95 a | 115 c | 158 c | |
Vs32972 | 44.67 e-k | 2.83 n-p | 3.00 g-i | 4.00 a-c | 0.57 bc | 7.11 a-f | 48.00 c-h | 15.10 c-f | 28 a | 95 a | 120 b | 162 b | |
Vs33456 | 43.43 e-k | 6.07 c-m | 4.86 f-i | 4.04 a-c | 0.64 bc | 6.39 b-i | 21.29 h-n | 5.30 g-n | 28 a | 95 a | 115 c | 158 c | |
Vs38517 | 34.67 f-k | 6.17 c-l | 3.33 f-i | 2.50 f-k | 0.53 bc | 4.72 g-n | 11.27 j-n | 3.05 h-n | 28 a | 95 a | 115 c | 158 c | |
Vs38523 | 43.33 e-k | 7.33 c-g | 3.67 f-i | 3.17 b-h | 0.60 bc | 5.28 f-m | 25.87 g-n | 7.17 f-n | 28 a | 95 a | 115 c | 158 c | |
Vs38526 | 86.17 b-d | 6.50 c-j | 6.33 f-i | 2.73 e-k | 0.40 bc | 6.18 b-j | 18.92 h-n | 5.50 g-n | 28 a | 95 a | 115 c | 158 c | |
Vs38527 | 90.00 bc | 9.83 b | 3.00 g-i | 4.53 a | 0.60 bc | 7.56 a-e | 20.41 h-n | 3.91 h-n | 28 a | 95 a | 115 c | 162 b | |
Vs38528 | 53.40 c-k | 8.00 b-e | 6.40 d-f | 3.63 a-f | 0.63 bc | 5.75 b-k | 34.49 e-l | 8.91 e-l | 28 a | 95 a | 115 c | 158 c | |
Vs38531 | 46.00 e-k | 6.40 c-k | 5.20 f-i | 2.84 c-j | 0.60 bc | 4.68 h-n | 13.00 i-n | 3.95 h-n | 28 a | 95 a | 115 c | 158 c | |
Vs38532 | 29.80 g-k | 7.60 b-f | 4.80 f-i | 2.00 f-k | 0.64 bc | 3.27 l-p | 8.35 k-n | 2.00 i-n | 28 a | 95 a | 115 c | 162 b | |
Vs38533 | 33.30 f-k | 4.74 f-p | 3.40 f-i | 3.66 a-f | 0.62 bc | 5.93 b-k | 10.54 j-n | 2.29 i-n | 28 a | 95 a | 115 c | 158 c | |
Vs38536 | 36.33 f-k | 7.00 c-h | 3.67 f-i | 2.23 g-k | 0.50 bc | 4.47 h-o | 3.04 mn | 0.83 k-n | 28 a | 95 a | 115 c | 158 c | |
Vs40310 | 33.33 f-k | 3.17 m-p | 3.67 f-i | 3.33 b-g | 0.50 bc | 6.67 a-h | 55.00 c-g | 15.77 c-e | 28 a | 95 a | 115 c | 158 c | |
Vs40315 | 38.33 f-k | 2.50 n-p | 3.67 f-i | 3.17 b-h | 0.50 bc | 6.33 b-i | 51.33 c-g | 14.63 c-f | 28 a | 95 a | 115 c | 158 c | |
Vs40326 | 44.67 e-k | 2.33 op | 4.33 f-i | 3.50 a-f | 0.50 bc | 7.00 a-f | 103.67 a | 36.33 a | 28 a | 95 a | 120 b | 162 b | |
Vs40334 | 31.33 f-k | 1.83 p | 4.67 f-i | 3.33 b-g | 0.43 bc | 7.83 a-c | 41.50 e-i | 8.93 e-l | 28 a | 95 a | 115 c | 158 c | |
Vs43100 | 22.00 j-k | 1.83 p | 4.33 f-i | 1.50 kl | 0.50 bc | 3.00 n-p | 9.57 j-n | 0.33 mn | 28 a | 95 a | 120 b | 162 b | |
Vsa38524 | 29.60 g-k | 7.70 b-f | 4.40 f-i | 3.16 b-h | 0.78 ab | 4.07 j-p | 7.18 l-n | 1.95 j-n | 28 a | 95 a | 115 c | 158 c | |
Vsa38525 | 50.44 d-k | 8.28 b-d | 5.56 f-h | 4.22 ab | 0.61 bc | 6.96 a-g | 19.92 h-n | 5.62 g-n | 28 a | 95 a | 115 c | 162 b | |
Vsa38530 | 65.33 b-h | 6.67 c-i | 2.00 i | 3.17 b-h | 0.53 bc | 5.69 c-k | 29.33 e-n | 10.98 d-h | 28 a | 95 a | 115 c | 162 b | |
Vsa38534 | 19.00 k | 5.50 d-n | 5.00 f-i | 3.25 b-h | 0.50 bc | 6.50 b-i | 26.75 f-n | 5.80 g-n | 28 a | 95 a | 115 c | 162 b | |
Vsa38535 | 36.50 f-k | 6.50 c-j | 2.50 hi | 2.75 d-k | 0.65 bc | 4.29 i-o | 4.55 l-n | 1.25 k-n | 28 a | 95 a | 115 c | 158 c | |
Vsa38537 | 43.67 e-k | 8.00 b-e | 5.67 f-h | 2.93 c-i | 0.60 bc | 4.89 f-n | 4.70 l-n | 1.05 k-n | 28 a | 95 a | 115 c | 162 b | |
Vsa4740 | 49.67 d-k | 3.50 k-p | 5.00 f-i | 2.00 h-l | 0.50 bc | 4.00 j-p | 45.00 f-h | 9.33 e-k | 21 b | 90 b | 107 d | 150 d | |
Vsa7243 | 54.33 c-k | 5.17 e-o | 4.33 f-i | 2.67 e-k | 0.50 bc | 5.33 e-m | 18.67 h-n | 5.67 g-n | 21 b | 90 b | 107 d | 150 d | |
Vsa38529 | 30 f-k | 4 i-p | 3 g-i | 0.6 m | 0.3 c | 2.00 p | 3.10 mn | 0.60 l-n | 28 a | 95 a | 115 c | 158 c | |
Vsc34295 | 85.13 b-d | 8.69 bc | 8.75 c-e | 1.06 l | 0.29 c | 3.75 k-p | 27.92 e-n | 6.92 f-n | 28 a | 95 a | 115 c | 158 c | |
Vss1862 | 44.67 e-k | 4.33 h-p | 4.33 f-i | 2.00 | 0.50 bc | 4.00 j-p | 31.67 e-m | 9.00 e-l | 21 b | 90 b | 107 d | 150 d | |
Vss24631 | 51.00 d-k | 2.33 op | 5.00 f-i | 1.83 i-l | 0.50 bc | 3.67 k-p | 56.67 b-f | 11.43 d-h | 21 b | 90 b | 125 a | 167 a | |
Vss29802 | 67.33 b-g | 3.50 k-p | 4.33 f-i | 2.50 h-l | 0.50 bc | 5.00 f-n | 41.50 e-i | 8.93 e-l | 28 a | 95 a | 107 d | 150 d | |
Vss32900 | 55.67 c-k | 3.00 n-p | 4.67 f-i | 1.67 j-l | 0.50 bc | 3.33 l-p | 34.50 e-l | 7.93 e-n | 28 a | 95 a | 107 d | 150 d | |
Vv315 | 55.00 c-k | 2.83 n-p | 4.33 f-i | 2.00 h-l | 0.50 bc | 4.00 j-p | 53.83 c-g | 13.08 c-g | 21 b | 90 b | 107 d | 150 d | |
Vv322 | 100.33 b | 4.00 i-p | 4.33 f-i | 2.50 f-k | 0.50 bc | 5.00 f-n | 108.33 a | 38.60 a | 28 a | 95 a | 115 c | 158 c | |
Vv6268 | 47.67 e-k | 3.17 m-p | 4.67 f-i | 1.67 j-l | 0.53 bc | 3.17 m-p | 83.50 ab | 19.73 bc | 21 b | 90 b | 107 d | 150 d | |
Vv14561 | 58.33 c-j | 4.50 g-p | 4.67 f-i | 2.67 e-k | 0.50 bc | 5.33 f-n | 57.71 b-e | 10.83 d-i | 28 a | 95 a | 115 c | 158 c | |
Vv28061 | 51.67 d-k | 2.00 p | 4.33 f-i | 2.00 h-l | 0.50 bc | 4.00 j-p | 18.00 h-n | 3.73 h-n | 28 a | 95 a | 115 c | 158 c | |
Vv34212 | 68.33 b-f | 3.67 j-p | 4.67 f-i | 2.17 g-l | 0.53 bc | 4.06 j-p | 39.33 e-i | 7.80 e-n | 28 a | 95 a | 115 c | 158 c |
Different letters indicate significant differences among different populations for the same species. P <0.05.
V. michauxii (Vmi), V. michauxii var. stenophylla (Vmis), V. monantha (Vmo), V.narbonensis (Vn), V. sativa (Vs), V. sativa var. angustifollia (Vsa), V. sativa var. cordata (Vsc), V. sativa var. sativa (Vss), V. villosa (Vv).
The results of phenology traits showed that all of the populations based on day to sprout and days to first flowering traits were divided into two groups (a and b). V. narbonesis (34878), V. monantha (32845) and two taxa of V. michauxii (Vmi2944 and Vmis37129) had the same value in day to sprout and days to first flowering traits, but populations V. sativa var.angustifollia (Vsa4740, Vsa7243), V. sativa var.stenophylla (Vss1862, Vss24631) and two populations of V. villosa (Vv315, Vv6268) with 21 and 90 d for day to sprout and first flowering were separated from the rest of populations by earlier germination and flowering. In days to total flowering and maturity traits, populations were divided as four groups (a, b, c and d). Days to total flowering as four groups:125a,120b,115c,107d and seed maturity:167a,162b,158c,150d. Populations in group d (107 and 150 d of flowering and seed maturation) had the shortest time required for full flowering and seed maturation. That is, they reached full flowering and seed maturity earlier than other populations. Populations of V. sativa var.angustifollia (Vsa4740, Vsa7243), V. sativa var.stenophylla (Vss1862, Vss29802, Vss32900) and V. villosa (Vv315, Vv6268), having the shortest day for full flowering and seed maturation (Table 4).
Analysis of the genetic correlations among the mentioned traits in the tested vetch populations revealed the existence of several significant positive coefficients (Table 5), namely between plant height with internode length (rgxy=0.43; P<0.01), stems number (rgxy=0.38; P<0.01) and pod length with internode length (rgxy=0.24; P<0.05), pod width (rgxy =0.23; P<0.05), day to sprout (rgxy=0.28; P<0.05), days to first flowering (rgxy=0.28; P<0.05) and days to maturity (rgxy=0.26; P<0.05), pod index with day to sprout (rgxy =0.23; P<0.05) and days to first flowering (rgxy=0.23; P<0.05). On the other hand, the relationship between pod width with pod index (rgxy =−0.26; P<0.05), biomass yield (rgxy =−0.35; P<0.01), and dry weigth (rgxy =−0.28; P<0.05), internode length with dry weigth (rgxy=−0.38; P<0.01) were negatively and significant.
Traits | Plant height |
internode length |
Stems number |
Pod length |
Pod width |
Pod index |
Biomass yield |
Dry weight |
Day to sprout |
Days to first flowering |
Days to total |
---|---|---|---|---|---|---|---|---|---|---|---|
Internode length | 0.43** | ||||||||||
Stems number | 0.38** | 0.11ns | |||||||||
Pod length | 0.13 ns | 0.24* | 0.03 ns | ||||||||
Pod width | 0.10 ns | 0.51 ns | -0.22 ns | 0.23* | |||||||
Pod index | 0.09 ns | -0.06 ns | 0.16 ns | 0.86 ns | -0.26* | ||||||
Biomass yield | 0.11 ns | -0.46 ns | 0.14 ns | -0.03 ns | -0.35** | 0.20 ns | |||||
Dry weight | 0.13 ns | -0.38** | 0.07 ns | 0.03 ns | -0.28* | 0.22 ns | 0.95 ns | ||||
Day to sprout | 0.04 ns | 0.19 ns | 0.06 ns | 0.28* | 0.11 ns | 0.23* | -0.16 ns | -0.09 ns | |||
Days to first flowering | 0.04 ns | 0.19 ns | 0.06 ns | 0.28* | 0.11 ns | 0.23* | -0.16 ns | -0.09 ns | 1 ns | ||
Days to total flowering | -0.08 ns | -0.03 ns | 0.05 ns | 0.17 ns | 0.03 ns | 0.15 ns | 0.04 ns | 0.07 ns | 0.46 ns | 0.46 ns | |
Days to maturity | -0.07 ns | 0.13 ns | 0.02 ns | 0.26* | 0.06 ns | 0.21 ns | -0.03 ns | 0.02 ns | 0.50 ns | 0.50 ns | 0.92 ns |
*, ** significant at 0.05 and 0.01 levels, respectively; ns not significant.
Two-dimensional principal component analysis showing the relationship among quantitative traits of studied populations is presented in Figure 1. Populations V. sativa var.angustifollia (4770, 7243), V. sativa var.sativa (1862), V. villosa (315, 6268) were separated partially by PC1; traits related to this separation are mainly phenology traits (day to spourat, days to first flowering, days to total flowering, days to maturity).
A cluster analysis of the tested Vicia spp. populations showed five main groups (Table 6 and Figure 2). Cluster G1 contained five populations, belonging to V.sativa var.angustifollia with two populations (7243, 4740), V. sativa var.sativa one population (1862) and V. villosa with two populations (315, 6268).They are characterized by the lowest values of phenology traits (day to spourat, days to first flowering, total flowering, and seed maturity). Cluster G2 contained 13 populations: 11 populations belonging to V. sativa (6646, 6681, 11761, 24062, 24069, 24074, 32972, 40310, 40315, 40326, 40334), population 38530 of V. sativa var.angustifollia and 322 of V. villosa. They are also characterized by the highest amount of vegetative, seed and yield traits compared to other populations. Cluster G3 included 16 popullations belonging to V. sativa (6654, 11760, 11762, 11771, 11772, 24076, 24084, 24097, 43100), population 38529 of V. sativa var.angustifollia, V. sativa var.sativa (24631, 29802, 32900), 28061, 34212 and 14561 of V. villosa, with high amount of vegetative traits were collected in one group. Cluster G4 contained seven populations: five belong to species V. sativa (11763, 11764, 11774, 38526, 38527), population 34295 of V. sativa var.cordata and V. monantha (32845). These were classified with the highest plant height, stems number and vegetative traits compared with other clusters. Cluster G5 was the largest one with 17 populations, nine from V. sativa (5321, 33456, 38517, 38523, 38528, 38531, 38532, 38533, 38536), five from V. sativa var.angustifollia (38524, 38525, 38534, 38535, 38537), V. michauxii (2944), V. michauxii var.stenophylla (37129) and V. narbonensis (34878).These were classified as highest vegetative and pod traits populations.
Groups | Plant height (cm) |
internode length (cm) |
Stems number |
Pod length (cm) |
Pod width (cm) |
Pod index |
Biomass yield (g) |
Dry weight (g) |
Day to sprout |
Days to first flowering |
Days to total flowering |
Days to maturity |
---|---|---|---|---|---|---|---|---|---|---|---|---|
G1 | 50.27 b | 3.80 c | 4.53 c | 2.07 c | 0.51 b | 4.10 d | 46.53 b | 11.36 b | 21.00 c | 90.00 c | 107.00 c | 150.00 c |
G2 | 50.97 b | 3.40 c | 4.36 c | 3.24 a | 0.50 b | 6.51 a | 63.15 a | 18.94 a | 28.00 a | 95.00 a | 116.15 a | 159.54 a |
G3 | 49.50 b | 3.43 c | 4.60 b | 2.04 c | 0.49 c | 4.14 d | 37.73 c | 8.20 c | 27.56 b | 94.69 b | 115.56 b | 158.31 b |
G4 | 88.76 a | 6.98 b | 9.25 a | 3.00 b | 0.48 c | 5.93 b | 30.92 c | 7.14 c | 28.00 a | 95.00 a | 115.71 b | 159.14 a |
G5 | 43.91 c | 7.12 a | 4.21 c | 3.01 b | 0.68 a | 4.72 c | 13.43 d | 3.44 d | 28.00 a | 95.00 a | 115.29 b | 159.18 a |
abc Different letters indicate significant differences among different populations for the same species. P<0.05.
The principal component analysis (PCA) of the 12 quantitative traits is summarized in Table 7. The first five PCs had eigenvalues >1 and they explained more than 80 % of the total variation for the vegetative and phenology traits. Day to sprout, days to first flowering, days to total flowering and days to maturity were loaded highly in PC1 and they accounted for 25.7 % of the total variation. In PC2, Biomass yield and dry weight accounted for 21 % of the total variation. In PC3, plant height and internode length accounted for 14.3 % of the total variation. PC4 contributed 11.2 % of the total traits variation in these populations with plant length and stems number loading highly. PC5 accounted for 9.8 % of total variation with length, width and pod index. Generally, for the 12 vegetative and phenology traits studied, PC1 and PC2 constituted more than 46 % of the total traits variation with most phenology traits and yield-related traits. This indicated that these traits can be used to classify the populations under study.
Variable | PC1 | PC2 | PC3 | PC4 | PC5 |
---|---|---|---|---|---|
Plant height | 0.058 | -0.029 | 0.466 | 0.441 | -0.267 |
Internode length | 0.208 | 0.302 | 0.428 | 0.055 | -0.242 |
Stems number | 0.050 | -0.107 | 0.176 | 0.668 | 0.075 |
Pod length | 0.310 | -0.154 | 0.389 | -0.244 | 0.378 |
Pod width | 0.141 | 0.303 | 0.243 | -0.301 | -0.357 |
Pod index | 0.230 | -0.332 | 0.265 | -0.084 | 0.533 |
Biomass yield | -0.144 | -0.53 | -0.004 | 0.062 | -0.236 |
Dry weight | -0.101 | -0.541 | 0.043 | -0.042 | -0.321 |
Day to sprout | 0.464 | -0.017 | -0.102 | 0.045 | -0.075 |
Days to first flowering | 0.464 | -0.017 | -0.102 | 0.045 | -0.075 |
Days to total flowering | 0.380 | -0.129 | -0.338 | 0.057 | -0.164 |
Days to maturity | 0.419 | -0.100 | -0.267 | 0.014 | -0.151 |
Eigenvalue | 3.340 | 2.723 | 1.856 | 1.452 | 1.279 |
Proportion | 0.257 | 0.210 | 0.143 | 0.112 | 0.098 |
Cumulative | 0.257 | 0.467 | 0.610 | 0.721 | 0.820 |
Discussion
In these study, 58 populations of Vicia spp. were investigated for genetic diversity based on morphological and phenology traits. Due to, genetic diversity analysis of germplasms using morphological traits is an initial step for crop improvement19-22. There was significant (P<0.01) genotypic variation among 58 germplasm accessions of Vicia spp. for all the measured vegetative and yield traits: plant length, internode length, stems number, pod length, pod width, pod index, biomass yield and dry weight.The estimates of genotypic variation and repeatability for these traits indicated the potential genetic variation available among the germplasm accessions within Vicia spp. investigated. Similar results were obtained by the Ebrahimi et al23 on plant and seed morphology traits of white Bean genotypes, Mikic et al12 on forage and seed yields of three lines of common vetch and Berhanu and Abera24 on forage yield of vetch species investigation.
A comparison between taxon (V. sativa: Vs, Vsa, Vsc and Vss, V. mchauxii: Vmi and Vmis, V. monantha: Vmo, V. narbonensis: Vn and V. villosa: Vv) showed V. monantha (Vmo) with high values of plant height, stems number, pod length and V. villosa with high values of biomass yield and dry weight. Berhanu and Abera24 showed that among the vetch species (V. sativa, V. villosa, V. dasycarpa, and V. bengalensis), V. dasycarpa and V. villosa were the best performing species for forage. Then the vetch species tested in the current study could be used for pasture expansion and forage production, in livestock exclusion areas, in forage strips, as an under-sowing with food crops, or as a backyard forage crop in the pasture of the country.
The populations demonstrated high variation in plant height, internode length, stem number, pod length, biomass yield and dry weight. Populations: Vmo32845, Vv322, Vmis37129 (for plant height), populations: Vmi37129, Vs38527, Vsc34295 (for internode length), populations: Vs11774,Vs11764,Vs11763 (for stems number), populations: Vs38527, Vmo32845, Vsa38525 (for pod length) and populations:Vs11761, Vs24062, Vs40326, Vv322, Vv6268 (for biomass yield and dry weight) showed the highest values of the mentioned traits. However, broadening the genetic base from diverse sources is recommended to include most of the genetic determinants of these traits25. This variability can be exploited in fodder breeding programs to select an adapted plant material for the arid and semi-arid areas26.
Phenology (earliness and lateness) of vetch species has a great effect on seed yield productivity. Late maturity for forage and seed was recorded at 125 and 167 d, respectively. This could be due to high and extended rainfall at the region of populations that encouraged vegetative growth and delayed forage and seed harvesting stages. The results indicated that for vetch populations tested, 107 to 125 and 150 to 167 d were required after the emergence of the seedlings for total flowering and seed maturity, respectively. On average, the difference in harvest forage and seed yield between populations are about 18 and 17 d. This indicates different responses of the tested populations for these important agronomic traits.
According to Getnet et al27, Vicia narbonensis and Vicia sativa are early maturing and Vicia villosa is late maturing species. But in this study two populations of V. villosa (315 and 6268) and four populations of V.sativa var.angustifollia (4740, 7243), V. sativa var.sativa (1862, 29802, 32900) with 107 and 150 d for flowering and seed maturity is recommended for seed production due to earliness, whereas late-maturing species like V. sativa var.sativa (24631) should not be advisable to grow for seed purpose.
There is a direct relationship between plant length with internode length and the number of stems, this indicates that tall plants produce long internodes and more number stems. Also, the length and width of the pod have a direct relationship with the number of days of sprout, flowering and seed maturation, which means long and wide pods are produced by late-flowering and seed maturation time. Since, in cereals, the correlation between grain yield and plant height is often negative, but in legumes, this correlation is often positive, because legumes have unlimited growth, therefore, with increasing height, more pods are produced, which has a positive effect on performance, so similar results were obtained in the traits of white Bean genotypes where high grain yields were strongly correlated to days to flowering and plant height23 and Lens spp.28.
In PCA, since the first component includes changes that are not explained by the second component and the two components are independent of each other, so the two components were intersected vertically and in the form of a biplot diagram to determine the diversity between different genotypes and determine the far and near genotypes to be used. Phenology traits (day to sprout, days to first flowering, days to total flowering, days to maturity) accounted for the variations recorded in the populations in PC1. On the other hand, yield traits (biomass yield and dry weight) accounted for the variation observed in the populations in PC2. The total cumulative variance in the first two PC was more than 46 %, indicating the high degree of diversity among the traits under study. Furthermore, the traits can be used as phenotypic traits in differentiating the populations. In plot PCA (Figure 1), populations, V. sativa var.angustifollia (Vsa7243, Vsa4740), V. sativa var.sativa (1862) and V. villosa (Vv315, Vv6268), separated from other populations and located on the left of X-axis by containing less of phenology traits (important in the first component). So, these populations recommend for areas with short growth periods. Populations V. sativa (40326) and V. villosa (322), for containing high value of biomass and dry weight, located on the bottom of Y-axis (negative effect of biomass and dry weight on the second component). As a result, two populations, V. sativa (40326) and V. villosa (322), produce forage yield more than other populations.
In the present study, the 58 populations of Vicia spp. were grouped into five clusters using 12 traits.The populations of cluster G1 are characterized by the lowest values of days to sprout, flowering, and seed maturity which are the candidate of further evaluations. Also, these populations had a shoter time for these traits. Members of G1 are similar to the dispersion of these populations in the PCA plot (Figure 1). It is interesting that the population from different climates like Shiraz clustered with populations from Karaj. This pattern of clustering indicates, the diversity of populations within these geographical areas and, the similarity of populations from different geographical areas.
These results agree with the report of Alemayehu and Becker29 in Brassica carinata. Cluster G2 contained 13 populations belonging to V. sativa and V. villosa species. These populations had a high value of seed, yield and phenology traits. Member of G2 due to having a long time for flowering and seed maturity, produce more seed and forage yield. This is the best factor, that can be used for livestock feeding. Cluster G3 contained mixed 16 populations of V. sativa and V. villosa. with lowest values of seed and forage yield gather together in a group, that they are not important inbreeding. Cluster G4 contained seven populations of V. sativa and V. monantha with high vegetative traits, that recommend livestock feeding and control of erosion. G5 group with 17 populations of V. sativa, V. michauxii and V. narbonensis were classified as later flowering and seed ripening and containing less amount of yield forage. These populations can be used for areas with a long growth time.
Finally in this study populations were located as five groups based on morphology and phenology traits. Members of each group are similar for mentioned traits and can be recommended for breeding programs. Also, the results indicated no relationship between studied traits and the origin of populations.
Conclusions and implications
The findings showed the high variation of morphology and yield traits in different species and populations of vetch. These differences are very important to select the type of companion crops and methods of integration to improve yields of both crops (food and forage) without significant effect of one on the other. Vicia sativa (Kermanshah, Javanrod) and V. villosa (Karaj) were superior in terms of fresh and dry forage yields. V.michauxii var.stenophylla (Qom), V. monantha (Kermanshah), V. sativa (Gilan, Astara), and V. villosa (Karaj), are recommended by having tall plant and big pods. However, more comprehensive studies and additional experiments are required to complete information for breeding programs.