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Abstract

Landraces are a valuable source of genetic variability for breeders to develop high-yielding lentil varieties. Apart from productivity, simultaneous breeding for lentil seed nutritional quality is of paramount importance for wider lentil consumption. This work examined the indirect effect of single plant selection for high yield on important seed quality traits within three Greek lentil landraces (“Elassona” (EL), “Lefkada” (L), and “Evros” (EV)). The breeding methodology applied was proved to help either maintain or improve such characteristics in the high-yielding second-cycle lines (SLs) selected. Compared to the parental landrace “Elassona”, the high-yielding lines showed increased crude fiber by 30–110%; the line 2-SL-EL-6 had higher starch content by 3.9% and reduced cooking time by 6.67 min, while the 2-SL-EL-10 line had higher crude fiber by 73%. In the case of “Lefkada”, the high-yielding lines selected maintained the protein content present in the parental landrace, apart from the 2-SL-L-1 where a decrease by 5% was recorded; however, most of them showed increased crude fiber (5.59–7.52%) in comparison with the parental landrace (4.65%). Finally, in relation to the “Evros” parental landrace, the 2-SL-EV-3 and 2-SL-EV-4 showed higher crude fiber and reduced cooking time. This study provides evidence that proper management of genetic variability could improve productivity without compromising or sometimes improving some seed quality traits.
agriculture
Article
Intense Breeding within Lentil Landraces for
High-Yielding Pure Lines Sustained the Seed
Quality Characteristics
Elissavet Ninou 1, * , Fokion Papathanasiou 2, Dimitrios N. Vlachostergios 1, Ioannis Mylonas 3,
Anastasia Kargiotidou 1, Chrysanthi Pankou 4, Ioannis Papadopoulos 2, Evangelia Sinapidou 5
and Ioannis Tokatlidis 5
1
Institute of Industrial and Forage Crops, Hellenic Agricultural Organization Demeter, 41335 Larissa, Greece
2Department of Agriculture, University of Western Macedonia, 53100 Florina, Greece
3Institute of Plant Breeding and Genetic Resources, Hellenic Agricultural Organization Demeter,
57001 Thermi, Greece
4School of Agriculture, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece
5Department of Agricultural Development, Democritus University of Thrace, 68200 Orestiada, Greece
*Correspondence: lisaninou@gmail.com
Received: 30 June 2019; Accepted: 6 August 2019; Published: 8 August 2019


Abstract:
Landraces are a valuable source of genetic variability for breeders to develop high-yielding
lentil varieties. Apart from productivity, simultaneous breeding for lentil seed nutritional quality is of
paramount importance for wider lentil consumption. This work examined the indirect eect of single
plant selection for high yield on important seed quality traits within three Greek lentil landraces
(“Elassona” (EL), “Lefkada” (L), and “Evros” (EV)). The breeding methodology applied was proved
to help either maintain or improve such characteristics in the high-yielding second-cycle lines (SLs)
selected. Compared to the parental landrace “Elassona”, the high-yielding lines showed increased
crude fiber by 30–110%; the line 2-SL-EL-6 had higher starch content by 3.9% and reduced cooking
time by 6.67 min, while the 2-SL-EL-10 line had higher crude fiber by 73%. In the case of “Lefkada”,
the high-yielding lines selected maintained the protein content present in the parental landrace, apart
from the 2-SL-L-1 where a decrease by 5% was recorded; however, most of them showed increased
crude fiber (5.59–7.52%) in comparison with the parental landrace (4.65%). Finally, in relation to
the “Evros” parental landrace, the 2-SL-EV-3 and 2-SL-EV-4 showed higher crude fiber and reduced
cooking time. This study provides evidence that proper management of genetic variability could
improve productivity without compromising or sometimes improving some seed quality traits.
Keywords:
lentil landrace; seed quality; physicochemical characteristics; cooking time; protein;
starch; crude fiber; breeding
1. Introduction
Lentil (Lens culinaris Medik.) is probably the oldest grain legume to be domesticated [
1
] and one
of the most important pulse crops worldwide due its nutritional characteristics. It is considered an
excellent source of complex carbohydrates, protein, minerals, vitamins, and dietary fibers [
2
,
3
]. Lentil
is a highly nutritious legume both as human food and animal feed, and the chemical composition of the
seeds is aected by genetic and environmental factors [
4
]. Despite its agronomic and nutritional value,
lentil seed production remained at low levels and attracted much less attention by plant breeders than
cereal grains until recently [
4
]. Currently, the extensive recognition of lentil’s health benefits resulted
in breeding varieties that are more productive and nutritious [4,5].
Agriculture 2019,9, 175; doi:10.3390/agriculture9080175 www.mdpi.com/journal/agriculture
Agriculture 2019,9, 175 2 of 13
It is widely accepted that genotype aects lentil chemical properties considerably [
3
,
6
]; thus,
to identify lentil genotype variability for seed quality traits would be valuable for the improvement of
lentil quality and the production of varieties with high nutrition value, such as high protein content [
7
,
8
].
Wang and Daun [
6
] reported significant variability in protein content ranging from 24.3% to 30.2%,
whereas Wang et al. [
9
], who found crude protein content values between 251.5 and 292.5 g/kg dry
matter, concluded that cultivar moreover had a considerable impact on other nutritious constituents
like starch, ash, and soluble dietary fiber. Starch content constitutes the highest proportion of the seed
in lentil and was recorded to range between 35 and 53% [
10
], 49 and 65% [
11
], and 41 and 49% [
12
],
while, in the case of ash content, values were recorded ranging from 2.13–3.42% [
13
] and 2.3–3.5% [
2
].
As for dietary fiber, which includes the plant-cell skeletal remains that are resistant to digestion [
14
],
Huisman and van der Poel [
15
] and Hulse [
16
] found that lentil decorticated seeds contained 0.9 g of
crude fiber/100 g, and other researchers reported values between 3.8% and 6% of dry seed weight [
13
].
Interestingly, Sulieman [
17
] reported that cultivars with high starch content generally had lower lipid
content. Regarding lipid content, Devos [
10
] identified a range from 1–2%, while, in other studies,
values were documented ranging from 0.5–2.8% and 1–1.3% [
2
]. More specifically for fat, according to
Adsule et al. [
18
] and Muehlbauer et al. [
19
], lentil seeds contain approximately 0.6 g of fat/100 g of
dried seeds, whereas Hulse [16] found concentrations of 1.8 g of fat/100 g of decorticated lentil seeds.
Apart from seed chemical composition, genetic variability was documented for physicochemical
characteristics, seed size, and seed processing (soaking, cooking, and dehulling). Cooking quality is
connected with cooking time, which aects nutrient and anti-nutrient contents [
2
,
9
,
13
]. The seed coat
of pulses is often indigestible and may have a bitter taste; thus, it could aect cooking quality and
eventually consumption [
3
]. Seed coat and hot water ability to penetrate the cotyledon are genetically
controlled and, thus, cause variability in cooking time in lentil varieties [
20
22
]. The cooking time of
Turkish lentil varieties was recorded between 15.2 and 23.9 min [
13
]; Jood et al. [
11
] reported values
ranging between 38 and 43 min, while Vandenberg [
23
] recorded cooking times varying from 15 to
20 min. Another parameter is water retention, i.e., the ability of food material to hold water against
gravity, defined as “water absorption” [
24
]. Lentil genotype aected seed hydration capacity that
ranged from 0.028–0.053 g/seed according to Özer and Kaya [
13
], while other researchers reported a
range of 0.019–0.023 g/per seed in dierent lentil varieties [
11
]. Finally, significant dierences among
lentil varieties were documented in physical properties and morphological characteristics, such as
1000-grain weight and size values [13,21].
Seed cooking quality is one of the most important factors for the utilization of lentil as food
because it is generally consumed in its cooked form. This characteristic is associated with the ease and
cost of food preparation [
3
]; thus, emphasis should be put on improving or maintaining it. Selection
for the improvement of seed quality characteristics is possible in lentil since genetic variability was
recorded for total starch, protein, 1000-seed weight, seed color, and other quality characteristics [
25
].
Landraces present a valuable gene pool for a breeder to develop elite lines and varieties [
26
,
27
].
Vlachostergios et al. [
28
] applied intense single-plant selection among widely spaced individual plants
in three lentil landraces aiming at second-generation sister lines of high yielding potential. When
evaluated at farming density, the derived lines had mean grain yields 8%, 10%, and 20% higher
compared to their respective ancestors [
28
]. The above findings support the view that the development
of pure line cultivars that fully meet the needs of sustainable agriculture is possible [
29
] and, at the
same time, ensures optimum use of resources across variable conditions [
30
]. The main criterion for
the selection within the Greek landraces was the yielding potential [
28
]; however, seed quality is an
important parameter for lentil consumption; thus, the eect of the breeding method applied to the seed
quality characteristics should be evaluated. The aim of the current study was to investigate the indirect
eect of intense breeding for high yield on seed quality characteristics, as well as level of variability for
these traits among the derived second-generation sister lines.
Agriculture 2019,9, 175 3 of 13
2. Materials and Methods
2.1. Genetic Material
For the study, 30 second-generation single-plant sister lines were used, obtained via the honeycomb
breeding methodology [
29
]. Selection was applied within three lentil landraces grown under a
nil-competition regime, i.e., individual plants were widely spaced to preclude any plant-to-plant
interference for inputs. In brief, within each landrace, selection of 30 out of 1000 initially established
plants led to first-generation sister lines. Then, evaluation of the first-generation sister lines at the
nil-competition regime at three locations followed by single-plant selection within the outstanding
lines led to the development of 20 second-generation sister lines, which were further evaluated
according to the nil-competition methodology at three locations. Finally, the most promising lines
were tested under the common farming density at five environments to verify their yield superiority
over their respective parental landrace. A detailed description of the breeding procedure is provided
by Vlachostergios et al. [28], as presented in Figure 1.
Agriculture 2019, 9, x FOR PEER REVIEW 3 of 14
2. Materials and Methods
2.1. Genetic Material
For the study, 30 second-generation single-plant sister lines were used, obtained via the
honeycomb breeding methodology [29]. Selection was applied within three lentil landraces grown
under a nil-competition regime, i.e., individual plants were widely spaced to preclude any plant-to-
plant interference for inputs. In brief, within each landrace, selection of 30 out of 1000 initially
established plants led to first-generation sister lines. Then, evaluation of the first-generation sister
lines at the nil-competition regime at three locations followed by single-plant selection within the
outstanding lines led to the development of 20 second-generation sister lines, which were further
evaluated according to the nil-competition methodology at three locations. Finally, the most
promising lines were tested under the common farming density at five environments to verify their
yield superiority over their respective parental landrace. A detailed description of the breeding
procedure is provided by Vlachostergios et al. [28], as presented in Figure 1.
Seed material for quality analysis for the present study originated from one of the above final
trials at farming conditions. That trial was established at the farm of the Fodder Crops and Pastures
Institute in Larissa (39°36’ north (N), 22°25’ east (E), 74 meters above sea level (masl)) during 2015
under rain-fed conditions without any application of chemicals and fertilizers, while weeds were
handled by hand-weeding as described by Vlachostergios et al. [28]. The experimental design was a
randomized complete block design with three replications. The second-generation lines included in
the trial were (i) 10 from the landrace “Elassona” (EL), coded as “2-SL-EL”, (ii) 10 from the landrace
“Lefkada” (L) coded as “2-SL-L”, and (iii) seven from the landrace “Evros” (EV) coded as “2-SL-EV”,
whereas (iv) the three parental landraces (PL) were also included.
Figure 1. A line diagram of the breeding scheme including single-plant selection at the nil-competition
regime within the parental landrace (PL) and the sister lines (SLs), with final SL evaluation at a dense
stand (adapted from Vlachostergios et al. [28]). The seed material studied in the current work
originated from one of the above final trials at a dense stand.
Figure 1.
A line diagram of the breeding scheme including single-plant selection at the nil-competition
regime within the parental landrace (PL) and the sister lines (SLs), with final SL evaluation at a dense
stand (adapted from Vlachostergios et al. [
28
]). The seed material studied in the current work originated
from one of the above final trials at a dense stand.
Seed material for quality analysis for the present study originated from one of the above final
trials at farming conditions. That trial was established at the farm of the Fodder Crops and Pastures
Institute in Larissa (39
36’ north (N), 22
25’ east (E), 74 meters above sea level (masl)) during 2015
under rain-fed conditions without any application of chemicals and fertilizers, while weeds were
handled by hand-weeding as described by Vlachostergios et al. [
28
]. The experimental design was a
randomized complete block design with three replications. The second-generation lines included in
the trial were (i) 10 from the landrace “Elassona” (EL), coded as “2-SL-EL”, (ii) 10 from the landrace
“Lefkada” (L) coded as “2-SL-L”, and (iii) seven from the landrace “Evros” (EV) coded as “2-SL-EV”,
whereas (iv) the three parental landraces (PL) were also included.
Agriculture 2019,9, 175 4 of 13
2.2. Agronomic and Morphological Seed Characteristics
The 1000-seed weight in grams was determined by counting 200 lentil seeds using an electronic
seed counter and scales. The results are expressed as the mean of duplicate measurements. Seed
width and thickness were measured in mm for 20 seeds per plot. The seed yield as presented by
Vlachostergios et al. [28] is provided in relation with the seed quality traits.
2.3. Seed Coat Color
The seed coat color of intact lentil seeds was recorded by a Minolta CR-410 chroma meter (Minolta,
Osaka, Japan) using the granular material attachment CR-A50. Data were collected for L* =lightness
ranging from 0 (black) to 100 (white), a* =redness/greenness, b* =yellowness/blueness, C* =chroma,
and H =hue. A white porcelain reference plate (Y=93.66, X=0.3150, and y=0.3217) supplied with
the instrument was used for calibration. All color parameters for each sample were the instrument
average of three independent measurements.
2.4. Physicochemical Characteristics
Seed coat percentage was determined for 20 seeds per plot as the weight ratio between coat and
cotyledons expressed in percentage after removing the seed coat from the cotyledons, having soaked
and kept the seeds for 24 h at 105 C.
The hydration increase of lentil seeds was calculated as the percentage increase in mass of lentils
soaked in distilled water for 12 and 24 h. Hydration capacity expressed as hydration capacity per seed
was determined by dividing the mass gained by the seeds either in 12 or 24 h by the number of seeds
present in the sample [31]. All tests were carried out in duplicate.
Cooking time was estimated according to the method described by Iliadis [
32
]. Twenty grams of
lentil seeds from each plot were added in 200 mL of distilled water in 250-mL conical flasks placed
in a 100
C water bath. After 20 min of initial cooking, samples of 10 seeds were taken from each
flask at 5-min intervals. Needle intrusion depth was measured using a penetrometer (Sur PNR-6,
Berlin Germany) for loading of 50 g and gravity of 0.2 sec. Seeds were considered cooked when the
penetration value was 4 mm.
2.5. Nutritional Quality Seed Traits
The nutritional quality traits were determined in duplicate finely ground samples. Seeds were
ground in a Cyclotec mill to pass a 1.0-mm screen. The mineral ash percentage calculated on dry weight
basis, total lipids, crude fiber, and total protein content (measured with the Kjeldhal method,
N×6.25
)
were determined using ocial methods [33]. Carbohydrates were determined by the dierence.
2.6. Statistical Analysis
Analysis of variance (ANOVA) was conducted for randomized complete block design. The
significance level of all hypotheses tested was pre-set at p<0.05, using the Tukey test (p<0.05). Pearson
correlation coecients were also calculated for all traits. All statistical analyses were performed using
the SPSS software package (ver. 18. SPSS Inc., Chicago, IL, USA).
3. Results
The recorded values of the seed quality characteristics for the three parental landraces and
second-generation high-yielding sister lines [
28
] revealed variability in several traits. The eect of
the selection for high seed yield on important seed quality characteristics on each of the three Greek
landraces is present below in detail.
Agriculture 2019,9, 175 5 of 13
3.1. “Elassona” Second-Generation Sister Lines
Concerning the EL lentil landrace, the protein, ash, lipids, 1000-seed weight, seed diameter and
thickness, seed coat percentage, and hydration index were not aected by the selection for high yield
(Tables 1and 2). Similarly, seed coat color parameters (L*, a*, b*, C*, and H) and hydration parameters
were not aected (data not shown). However, the crude fiber values of the selected genotypes diered
from the EL parental landrace and ranged from 3.21% to 6.94%. In five selections, significant variability
was found, and an increase from 30% to 110% was recorded in comparison with the parental landrace.
The high-yielding selection 6 (2-SL-EL-6) had similar crude fiber concentration with the original
genotype, whereas the high-yielding selection 10 (2-SL-EL-10) showed increased crude fiber by 73% in
comparison with the EL parental landrace (Table 1).
Table 1.
Seed yield and nutritional quality seed traits of the parental landrace “Elassona” (EL-PL) and
second-generation sister lines (SLs) originating from “Elassona” (EL).
Lentil
Genotypes
Seed Yield
(kg/ha)
Protein
(%)
Ash
(%)
Crude Fiber
(%)
Lipids
(%)
Starch
(%)
EL-PL 1124 28.09 2.65 3.26 1.06 53.58
2-SL-EL-1 1278 26.80 3.05 4.81 ** 0.77 53.12
2-SL-EL-2 1170 27.30 3.02 3.35 0.63 54.47
2-SL-EL-3 1038 27.49 3.02 4.69 ** 0.72 52.66
2-SL-EL-4 1196 26.88 2.88 6.94 ** 0.48 51.12 **
2-SL-EL-5 1225 28.05 3.10 3.93 * 0.83 52.86
2-SL-EL-6 1387 * 26.24 2.88 3.21 0.69 55.67 **
2-SL-EL-7 1059 27.85 2.92 3.62 0.83 53.29
2-SL-EL-8 1210 27.30 3.16 3.76 0.84 53.57
2-SL-EL-9 1265 27.28 3.11 3.92 0.79 53.55
2-SL-EL-10 1352 * 27.66 2.99 5.65 * 0.72 52.35 *
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
Table 2.
Agronomical, morphological, seed coat %, hydration index, and cooking time of the parental
landrace “Elassona” and second-generation sister lines originating from “Elassona”.
Lentil
Genotypes
1000-Seed
Weight (g)
Diameter
(mm)
Thickness
(mm)
Seed Coat
Percentage (%)
Hydration
Index (12 h)
Hydration
Index (24 h)
Cooking
Time (min)
EL-PL 32.90 4.57 2.51 8.01 0.83 0.90 41.67
2-SL-EL-1 35.13 4.79 2.51 7.73 0.83 0.93 33.33 **
2-SL-EL-2 40.33 5.00 2.56 7.77 0.84 0.94 38.33
2-SL-EL-3 37.17 4.79 2.64 7.89 0.83 0.92 36.67 *
2-SL-EL-4 36.20 4.80 2.55 7.85 0.84 0.94 43.33
2-SL-EL-5 29.07 4.28 2.50 7.99 0.85 0.96 31.67 **
2-SL-EL-6 34.33 4.78 2.57 7.89 0.83 0.93 35.00 **
2-SL-EL-7 36.20 4.77 2.53 7.40 0.86 0.95 35.00 **
2-SL-EL-8 36.03 4.73 2.58 8.02 0.77 0.91 43.33
2-SL-EL-9 34.70 4.59 2.59 8.00 0.86 0.93 35.00 **
2-SL-EL-10 34.30 4.43 2.56 7.52 0.82 0.91 36.67
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
The starch was also aected by the selection for high yield and ranged from 51.12% to 55.67%.
It is noteworthy that, in selection 6 (2-SL-EL-6), the starch content increased by 3.9%. However,
in two selections (2-SL-EL-4 and 2-SL-EL-10), inferior values were observed, and the remaining
high-yielding sister lines showed no significant dierences in starch concentration (Table 1). Cooking
time connected with the consumption value of lentils varied from 31.67 to 43.33 min (Table 2).
Remarkably, the high-seed-yielding selection 6 had a reduced cooking time by 6.67 min compared to
the EL parental landrace.
In general, the protein content and seed quality characteristics of the high-yielding lines selected
were not aected by the breeding methodology, since no dierences were recorded in comparison with
the parental landrace. However, the high-yielding selection 6 had higher starch content and shorter
cooking time, while the high-yielding selection 10 was richer in crude fiber (Tables 1and 2).
Agriculture 2019,9, 175 6 of 13
3.2. “Lefkada” Second-Generation Sister Lines
The L lentil landrace and the high-yielding second-generation sister lines showed variability for
protein (26.8%–28.3%), crude fiber (3.89%–7.52%), and starch content (48.55%–53.9%) (Table 3).
Table 3.
Seed yield and nutritional quality traits of the parental landrace “Lefkada” (L-PL) and
second-generation sister lines originating from “Lefkada”.
Lentil
Genotypes
Seed Yield
(kg/ha)
Protein
(%)
Ash
(%)
Crude fiber
(%)
Lipids
(%)
Starch
(%)
L-PL 1136 28.33 2.72 4.65 0.92 51.67
2-SL-L-1 1455 ** 26.82 * 3.07 5.66 ** 0.67 52.36
2-SL-L-2 1343 ** 28.90 3.13 6.16 ** 0.73 49.39 *
2-SL-L-3 1379 ** 28.30 3.24 * 5.05 0.88 51.48
2-SL-L-4 1395 ** 27.42 3.50 7.30 ** 0.68 49.65
2-SL-L-5 1297 27.03 * 2.95 5.39 * 0.53 * 52.89
2-SL-L-6 1446 ** 27.72 2.67 7.52 ** 0.58 50.09
2-SL-L-7 1278 28.10 2.76 5.59 * 0.93 51.17
2-SL-L-8 1259 27.43 2.80 3.89 * 0.70 53.90 *
2-SL-L-9 1357 ** 27.80 2.64 6.66 ** 0.74 48.55 **
2-SL-L-10 1507 ** 27.31 3.28 5.01 0.77 52.32
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
The selection for high yield in the L lentil landrace did not aect the protein content in the majority
of the high-yielding sister lines compared with the parental landrace. Among the high-yielding
selections, only 2-SL-L-1, which had the second highest yield (1455 kg/ha), showed a decrease in
protein content by 5% (Table 4). However, the selection for high yield was accompanied by an increase
in crude fiber for eight selections in comparison with the parental landrace (Table 3).
Table 4.
Agronomical, morphological, seed coat %, hydration index, and cooking time of the parental
landrace “Lefkada” and second-generation sister lines originating from “Lefkada”.
Lentil
Genotypes
1000-Seed
Weight (g)
Diameter
(mm)
Thickness
(mm)
Seed Coat
Percentage (%)
Hydration
Index (12 h)
Hydration
Index (24 h)
Cooking Time
(min)
L-PL 30.27 4.34 2.56 8.55 0.83 0.92 33.33
2-SL-L-1 37.90 4.72 2.62 7.70 0.85 0.94 33.33
2-SL-L-2 34.83 4.74 2.54 8.20 0.80 0.95 26.67
2-SL-L-3 33.20 4.54 2.53 8.10 0.84 0.95 28.33
2-SL-L-4 36.27 4.61 2.48 7.81 0.82 0.92 31.67
2-SL-L-5 36.30 4.78 2.60 7.96 0.83 0.93 30.00
2-SL-L-6 38.27 4.90 2.55 7.89 0.83 0.92 31.67
2-SL-L-7 32.17 4.37 2.52 7.98 0.83 0.94 26.67
2-SL-L-8 35.10 4.63 2.49 8.16 0.80 0.90 30.00
2-SL-L-9 36.20 4.80 2.54 8.15 0.88 0.97 30.00
2-SL-L-10 31.85 4.22 2.55 7.96 0.81 0.89 35.00
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
Regarding starch content, the selection 2-SL-L-8 had higher starch content in comparison with the
parental landrace, although it was not dierentiated for seed yield (Table 3). The selections 2-SL-L-2
and 2-SL-L-9 showed reduced values by ~5%, while the other five high-yielding selections did not
dier for starch content (Table 3). The breeding scheme applied aimed at seed yield increase, and seven
out of the total 10 second-generation sister lines had higher yield. At the same time, the methodology
based on a single-plant selection sustained the other seed quality characteristics in the “Lefkada”
landrace. The 1000-seed weight, seed diameter and thickness, seed coat percentage, hydration index,
and cooking time remained constant in comparison with the “Lefkada” parental landrace (Table 4).
Also, the seed coat color parameters (L*, a*, b*, C*, and H) and the hydration increases, coecients,
and capacities were not altered (data not shown).
Agriculture 2019,9, 175 7 of 13
3.3. “Evros” Second-Generation Sister Lines
Concerning the EV lentil landrace, the protein, ash, lipids, 1000-seed weight, seed diameter and
thickness, seed coat percentage, and hydration index were not affected by the selection for high yield
(Tables 5and 6). Also, no effect on seed coat color parameters (L*, a*, b*, C*, and H) and hydration
increases, coefficients, and capacities was found (data not shown). Nevertheless, the seed characteristics
of the EV second-generation sister lines showed variability for crude fiber, starch content, and cooking
time ranging from 3.12% to 5.62%, 49.93% to 54.06%, and 31.67 to 41.67 min, respectively (Table 5).
Noticeably, six sister lines selected had increased crude fiber in comparison with the parental landrace
“Evros”, and two of them (2-SL-EV-3 and 2-SL-EV-4) had significantly higher seed yield (Table 5). The
starch content was decreased by 4.7% in the high-seed-yielding selection 2-SL-EV-3, while it was not
altered in the high-seed-yielding selection 2-SL-EV-4 compared to the parental landrace EV. Three
second-generation sister lines, including the two highest-yielding 2-SL-EV-3 and 2-SL-EV-4 selected lines,
showed a highly significant lower cooking time by almost 10 min compared to the original genotype.
Table 5.
Seed yield and nutritional quality traits of the parental landrace “Evros” (EV-PL) and
second-generation sister lines originating from “Evros”.
Lentil
Genotypes
Seed Yield
(kg/ha)
Protein
(%)
Ash
(%)
Crude Fiber
(%)
Lipids
(%)
Starch
(%)
EV-PL 1100 28.19 2.62 3.12 0.86 53.57
2-SL-EV-1 992 27.24 2.62 3.82 0.93 54.06
2-SL-EV-2 1194 29.44 2.64 5.62 ** 0.85 49.93 **
2-SL-EV-3 1455 ** 29.02 2.67 4.86 ** 0.96 51.07 *
2-SL-EV-4 1412 ** 27.64 2.74 4.91 ** 0.88 52.37
2-SL-EV-5 1188 27.37 2.51 4.08 * 1.11 53.46
2-SL-EV-6 1048 27.33 2.50 5.25 ** 0.89 52.52
2-SL-EV-7 1187 27.94 2.61 4.41 ** 0.90 52.61
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
Table 6.
Agronomical, morphological, seed coat %, hydration index, and cooking time of the parental
landrace “Evros” and second-generation sister lines originating from “Evros”.
Lentil
Genotypes
1000-Seed
Weight (g)
Diameter
(mm)
Thickness
(mm)
Seed Coat
Percentage (%)
Hydration
Index (12 h)
Hydration
Index (24 h)
Cooking
Time (min)
EV-PL 35.40 4.62 2.54 7.94 0.85 0.95 41.67
2-SL-EV-1 39.37 5.03 2.56 7.88 0.86 0.94 40.00
2-SL-EV-2 36.73 4.84 2.58 7.73 0.86 0.95 31.67 **
2-SL-EV-3 34.40 4.53 2.58 7.72 0.84 0.93 31.67 **
2-SL-EV-4 33.17 4.33 2.59 7.60 0.81 0.92 31.67 **
2-SL-EV-5 35.27 4.80 2.51 8.04 0.84 0.94 40.00
2-SL-EV-6 36.57 4.79 2.62 7.98 0.82 0.92 36.67
2-SL-EV-7 35.67 4.62 2.54 8.13 0.85 0.94 35.00
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
The application of the breeding scheme as described by Vlachostergios et al. [
28
] for two
selection cycles allowed the selection for higher yield without compromising important seed quality
characteristics for the majority of the genotypes selected compared to their respective parental landraces.
The variation and descriptive statistics for important parameters like nutritional quality traits (Table S1,
Supplementary Materials), agronomical, morphological, and seed coat characteristics (Table S2,
Supplementary Materials), and hydration parameters and cooking time (Table S3, Supplementary
Materials) were not dierentiated in comparison with the parental landraces.
Finally, the correlation between agronomical, morphological, physicochemical, and nutritional
quality traits and seed coat characteristics was estimated. A negative correlation was found between
starch content and crude fiber (r
st.-cf
=
0.856 **), whereas seed diameter and 1000-seed weight were
positively correlated (r
sd-1000sw
=0.881 **) (Table 7). Also, a positive correlation was found between
L* (lightness) and 1000-seed weight, b* (yellowness/blueness), C* (chroma), H (hue) (r
L-1000 SW
=
0.690 **, r
L-b
=0.855 **, r
L-C
=0.851 **, and r
L-H
=0.738 **, respectively), between H (hue) and b*
(yellowness/blueness) and C* (chroma) (r
H-b
=0.879 ** and r
H-C
=0.788 **, respectively), and between
C* (chroma) and b* (yellowness/blueness) (rC-b =0.985 **).
Agriculture 2019,9, 175 8 of 13
Table 7. Correlation between agronomical, morphological, physicochemical nutritional quality traits and seed coat characteristics.
Yield Protein Ash Crude
Fiber Lipids Starch 1000-Seed
Weight Diameter Thickness L* a* b* C* H Seed
Coat
Hydration
Increase (24 h)
Cooking
Time
Yield 1 0.005 0.359 0.468 ** 0.318 0.377
*0.180 0.326 0.024 0.052 0.214 0.314 0.290 0.281 0.145 0.129 0.541 **
Protein 1 0.203 0.107 0.425* 0.538
** 0.307 0.313 0.132 0.204 0.220 0.048 0.012 0.217 0.161 0.226 0.349
Ash 1 0.163 0.446
*0.064 0.062 0.148 0.091 0.080 0.010 0.161 0.175 0.099 0.145 0.050 0.211
Crude fiber 1 0.467
**
0.856
** 0.141 0.036 0.019 0.041 0.286 0.262 0.191 0.384 * 0.034 0.095 0.422 *
Lipids 1 0.151 0.388 * 0.303 0.190 0.243 0.088 0.133 0.124 0.122 0.189 0.020 0.132
Starch 1 0.044 0.102 0.089 0.127 0.360 0.273 0.188 0.452 * 0.126 0.285 0.517 **
1000-Seed
Weight 1 0.881 ** 0.301 0.690
** 0.017 0.583 ** 0.600 ** 0.458 * 0.336 0.075 0.291
Diameter 1 0.158 0.586
** 0.028 0.542 ** 0.547 ** 0.446 * 0.136 0.238 0.314
Thickness 1 0.032 0.040 0.042 0.054 0.006 0.179 0.163 0.123
L* 1 0.170 0.855 ** 0.851 ** 0.738 ** 0.503 ** 0.033 0.157
a* 1 0.221 0.064 0.605 ** 0.032 0.111 0.066
b* 1 0.985 ** 0.879 ** 439 * 0.032 0.423 *
C* 1 0.788 ** 451 * 0.016 0.417 *
H 1 378 * 0.021 0.394 *
Seed Coat 1 0.015 0.134
Hydration
Increase (24 h) 10.225
Cooking time 1
*, ** Significant dierences at the 0.05 and 0.01 levels of probability.
Agriculture 2019,9, 175 9 of 13
4. Discussion
Legumes could provide a sustainable solution for food production in terms of protein security.
Concerning nutrition, landraces have higher seed protein content than modern varieties, as recorded
in common bean [
34
]. In this sense, lentil landraces of good quality and high nutritional value that are
adapted to local environmental conditions present a valuable source of genetic variability [
28
]. In this
work, the selection of second-generation sister lines employed single-plant yield as the main selection
criterion [
28
] without focusing on quality characteristics as a direct selection criterion. The scope of
studying multiple quality characteristics was to investigate the indirect eect of this breeding scheme
on seed quality. Variability regarding important seed quality traits and physicochemical characteristics
was recorded for the three Greek lentil landraces and the second-generation sister lines examined.
The main significant dierences were identified in protein, crude fiber, starch content, and cooking
time. Remarkably, some high-yielding selected lines displayed unaltered protein content and other
important quality characteristics.
Iqbal et al. [
35
] stated that a major challenge in legume breeding is to increase productivity and
nutritional quality; however, they recognized that quality evaluation requires screening both in the
field and laboratory; thus, it is a time- and resource-consuming selection process. This method of
single-plant selection for high seed yield under a nil-competition regime presents an approach that
could minimize the time and resources required to improve productivity and maintain nutritional
seed quality. Interestingly, the same methodology was also recommended as an eective agronomic
practice to improve the sanitary status of lentil landraces seed stock during the seed propagation
process, especially in case of seed-borne viruses that are transmitted by seed in high rate [36].
The protein concentration measured in the present study ranged from 26% to 28%, which agrees
with the values reported by Wang and Daun [
6
], who calculated a mean level of 27.2% in protein
content. Likewise, protein contents reported in lentils were around 25% [
37
], between 24.3% and
30.2% [
12
], and between 22.1 and 27.4% [
38
]. In this work, the protein content was unaltered in
comparison with the parental landraces, even when the seed yield increased. Thus, direct selection
for high yield in the EL landrace helped to maintain the seed protein content. A similar picture was
drawn in the case of all the L high-yielding sister lines selected, apart from the 2-SL-L-1 that showed
a decrease by 5%. This was also the case for the protein content of the EV second-generation sister
lines that did not dier compared to the parental landrace, even for the two selections with the highest
yield (2-SL-EV-3 and 2-SL-EV-4). Such results are particularly promising since several researchers
recorded a negative correlation between yield and protein content, which can be alleviated by breeding
for both traits since sucient variation for these coexists in legumes [
39
]. Iqbal et al. [
35
] suggested
that an approach to exploit genotypic variability within legume populations is to select for increased
yield and simultaneously aim to maintain a constant protein level. The breeding scheme of this study
(Figure 1) [
28
] supplemented with the evaluation of quality traits could be proposed in alignment with
the above recommendation.
In terms of crude fiber, it ranged from 3.12% to 7.52% for the three landraces and their
second-generation sister lines studied; such levels are in agreement with those reported for Turkish
lentil varieties (between 3.8% and 6%) [
13
]. In this study, the selection for high yield was accompanied
by stable or increased crude fiber content in some selections originating from EL, L, and EV landraces.
The crude fiber content increased (2-SL-EL-10) or remained stable (2-SL-EL-6) in two high-yielding
selections from the EL landrace, while the crude fiber for the eight L selections increased. Dierences
in fiber content among lentil cultivars exist due to genetic variability, suggesting a rationale for
cultivar-based food labeling [40].
Regarding the starch content measurements in this study, a range from 48.55% to 55.67% was
close to levels previously reported [
38
]; this parameter was aected in comparison with the parental
landraces in some high-yielding selections of the EL, L, and EV landraces. Thus, the starch content in
the two high-yielding EL selections was reduced (2-SL-L-2 and 2-SL-L-9), while the starch content of
the EV selections decreased or remained constant in comparison with the EV parental landrace.
Agriculture 2019,9, 175 10 of 13
Lentil genetic resources display significant variability for seed quality characteristics such as color,
ranging from yellow to red-orange to green, brown, and black, as well as seed coat and cotyledon
color [
41
]. Other quality traits that may be aected by cultivar are cooking quality, seed size, uniformity,
and absence of split and discolored seeds [
42
]. We found that the intense breeding applied in three lentil
landraces not only resulted in maintaining color and seed size unaltered compared with the parental
landraces, but also resulted in improving cooking time in some of the lines selected. Jood et al. [
11
]
reported that lentil cultivars diered in seed hydration capacity, and flat seeds with large surface area
absorbed more water in comparison with those with narrow surface. A similar connection with seed
size and water absorbance was identified also by Özer and Kaya [
13
], since flat lentil varieties with
a large surface area revealed a more swollen appearance after water treatment, and varieties with
narrow surface area absorbed less water per seed. In the current work, no dierences were recorded
for hydration capacity, seed size, and seed color, apart from cooking time which decreased, especially
in the “Elassona” and “Evros” high-yielding selected lines by almost 6 and 10 min, respectively.
Several important qualitative characteristics examined like average seed weight, protein content,
and cooking quality showed high broad-sense heritability [
21
]. Plant breeders are interested mainly in
traits having high heritability since they result in greater advance under selection. The values reported
for seed weight of lentil were 62.8 % [
43
], 87.0% [
44
], 91.0% [
45
], 98.0% [
21
,
46
], and 99.0% [
47
]. High
broad-sense heritability was also found for protein content with a value of 84.0% [
47
] and cooking
quality with 82.0% [
21
] and 98.0% [
47
]. Finally, moderate to high broad-sense heritability was found for
lentil grain yield as reported by several researchers, ranging from 41.0% [
45
] to 48.0% [
44
], 59.7% [
43
],
83.9% [46], and 96.0% [47].
Several researchers reported correlations between dierent seed quality characteristics in lentil
genotypes. In this work, a negative correlation was found between starch content and crude fiber.
A negative correlation was identified between starch and protein content [
6
,
25
], and also 1000-seed
weight and protein concentration [
25
]. Similarly, a negative correlation was reported for seed yield and
protein content, whereas cooking time and seed weight were positively correlated [
21
]. Seed yield was
previously positively correlated to both cooking time and seed size, which aects cooking quality [
21
],
a sign that seed size might be used to predict cooking time [
47
]. However, in this work, no correlation
was found between cooking time and seed size, supporting the fact that the above might not be always
be the case. A positive correlation was found between the 1000-seed weight and seed starch, total
ranose family oligosaccharides, and sucrose, while there was no correlation between the seed coat
color and protein concentration [
48
]. Finally, in this work, while there was no correlation between the
protein content and seed color, the seed diameter and 1000-seed weight were positively correlated,
showing that the increase in seed weight is connected with an increase in seed diameter rather than
its thickness.
Breeding strategies include the development of more productive varieties with favorable quality
characteristics. In the case of lentil, research is underway for the development of improved plant types
with high yielding ability and resistance to biotic and abiotic stress, while better nutrition quality is
also a major breeding objective. So far, however, studies often focused on either yield gain or quality
improvement rather than the influence of breeding techniques on such parameters combined. The
procedure presented here provides the tools to exploit the natural genetic variability within landraces
and develop, in a short time, pure-line varieties adaptable to a wide range of conditions that moreover
display desirable characteristics [
28
]. It is recognized that yield is negatively correlated with protein
content, and it is important to understand the relationship among dierent quality parameters for the
selection of new cultivars with favorable quality characteristics [
25
]. There is evidence supported by
this study that the proper management of genetic variability shows potential for the simultaneous
increase of productivity and seed quality characteristics. There is need for more in-depth studies
regarding the nutritional quality of this low-cost protein source and the influence of compounds such
as fiber and related substances [49].
Agriculture 2019,9, 175 11 of 13
Even though huge genetic variation facilitates the selection of superior genotypes, and selection
within a landrace is definitely one of the oldest plant breeding methods, the innovation brought about
by the method presented here is that it leads toward the identification of a dierent class of genotypes
in order to develop varieties in accordance with the main principles of sustainability. Based on the
well-established negative relationship between genotype yield and competitive ability, breeding in
conditions that ensure nil-competition instead of dense stand, and applying single-plant yield as the
main selection criterion puts additional emphasis on highlighting the “weak competitor of high plant
yield eciency” ideotype [
30
]. The hypothesis of this approach is that genotypes characterized as
“weak competitors” are exceptionally resilient to environmental forces that induce acquired intra-crop
variation and, thus, could be employed to optimize resource use at crop level, while, at the same time,
eciency in resource use at single-plant level promotes stability due to better overall results in case of
missing plants in the field [50].
5. Conclusions
The development of varieties combining high productivity and favorable nutritional characteristics
constitutes a challenge in lentil breeding for increased consumption and widespread cultivation. The
applied breeding scheme and selection for high yield based on plant performance at ultra-low density
ensured in two cycles of selection that the seed quality characteristics of the landraces parental
population remained unaltered. This could be an eective strategy to manipulate the existing genetic
variability for seed quality and develop varieties combining higher seed yield and quality. According
to the results of this work, the proposed breeding procedure favors the selection of high-yielding
ideotypes that sustain qualitative characteristics. This innovative approach in breeding methodology
provides evidence that breeding for high yield without degrading quality traits might be feasible.
Supplementary Materials:
The following are available online at http://www.mdpi.com/2077-0472/9/8/175/s1,
Table S1: Descriptive statistics of nutritional quality traits of second-generation sister lines originating from
“Elassona”, “Lefkada”, and “Evros”, Table S2: Descriptive statistics of agronomical, morphological, and seed coat
characteristics of second-generation sister lines originating from “Elassona”, “Lefkada”, and “Evros”, Table S3:
Descriptive statistics of hydration increases, coecients, capacities, and indices and cooking times of each parental
landrace and second-generation sister lines originating from their respective parental landrace.
Author Contributions:
Conceptualization, I.T.; data curation, F.P. and I.M.; formal analysis, I.M.; funding
acquisition, I.T.; investigation, E.N.; methodology, F.P., D.N.V., and I.P.; project administration, I.T.; resources, E.N.,
A.K., C.P., and I.P.; writing—original draft, E.N., F.P., D.N.V., I.M., E.S., and I.T.; writing—review and editing, E.N.,
F.P., D.N.V., I.M., A.K., C.P., I.P., E.S., and I.T.
Funding: Research co-financed by the European Union (European Social Fund) and Greek national funds in the
framework of the project “THALIS—Democritus University of Thrace—Selection for enhanced yield and tolerance
to viral and vascular diseases within lentil landraces” through the Operational Program of the NSRF (National
Strategic Reference Framework) “Education and lifelong learning investing in knowledge society”.
Conflicts of Interest: The authors declare no conflict of interest.
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article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).
... The demand for human consumption of plant-based protein will continue to grow in the future (Pihlanto et al., 2017). Previous studies demonstrated that protein concentration in lentil varies by cultivar (Ghumman et al., 2016;Ninou et al., 2019;Nosworthy et al., 2018;Wang & Daun, 2006), climate (Erskine et al., 1985;Fatima et al., 2018;Sharaan et al., 2003), soil (Huang et al., 2016;Zeidan, 2007), and agricultural practices (Gill, 2013;Mandal et al., 2018). Wang and Daun (2006) found that protein concentration in lentil varied between 243 and 302 g kg −1 dry matter (DM), whereas in another study protein concentration ranged from 252 to 293 g kg −1 DM (Wang et al., 2008). ...
... The broad-sense heritability was estimated at .62 based on the four cultivars grown in five locations for 3 yr, indicating breeding cultivars for higher protein concentration is possible. Other studies have also reported significant differences in protein concentration amongst cultivars (Ghumman et al., 2016;Ninou et al., 2019;Nosworthy et al., 2018;Subedi et al., 2020;Tao et al., 2017;Wang & Daun, 2006). ...
Article
Lentil (Lens culinaris Medik.) is an important source of protein, starch, and mineral nutrients in many parts of the world. However, the impact of environment and cultivar on the enrichment of nutrients including protein, starch, and minerals is not well understood in important lentil production regions, including the U.S. Northern Great Plains. Four lentil cultivars (Avondale, CDC Richlea, CDC Maxim, and CDC Imvincible) varying in color, seed size and maturity were evaluated at five Montana locations with diverse climatic and soil conditions over three years. Significant cultivar, location, and year effects were found for yield, protein, starch, and minerals. Grain protein concentration was the highest at Moccasin (262 g kg–1) and lowest at Richland (246 g kg–1), while starch concentration was the highest at Richland (455 g kg–1) and lowest at Moccasin (441 g kg–1). Among cultivars, Avondale was the top yielding cultivar (1965 kg ha–1) and adaptable to most of the environments; CDC Imvincible was the top protein producer (265 g kg–1); and CDC Richlea is the leading starch producer (456 g kg–1). Grain protein concentration was negatively correlated with starch. Lentil grains varied in macro‐ and micro‐nutrient concentrations across locations, with the northcentral Montana region producing 10–20 times greater Se concentration than other locations. CDC Maxim had the highest Fe (62.1 mg kg–1) and Zn (31.5 mg kg–1) concentrations. Seed protein concentration was positively correlated with P, S, Cu, and B. Seed starch is positively correlated with Mg and Mn. Results suggest that plant breeding and production site selection could enrich lentil nutrient concentrations to help combat malnutrition in the world. This article is protected by copyright. All rights reserved Lentil grain produced at different locations showed significant differences in nutrient concentrations. Different cultivars showed different capacities of enriching protein and minerals Protein is negatively correlated with starch and positively correlated with certain mineral elements Plant breeding and production site selection could enhance lentil nutrient concentrations.
... Galiotou-Panayotou et al. (66) stated that a decrease in phytic acid from 10.4 to 7.2 mg g −1 induced a 42% increase in hardness. This suggests that low cookability of lentil seed grown under high temperature was due to the high level of phytic acid.Cooking time showed no significant correlation with 100-seed weight under both conditions, which was in accordance with finding reported by Ninou et al. (67). However, significant positive correlation has been reported in lentil (68) and common bean (69). ...
... Correlation analysis also showed that cooking time was positively correlated with seed thickness under normal conditions. In contrast, a nonsignificant correlation between cooking time and seed thickness has been demonstrated in lentil (67). Moreover, this seed shape parameter also showed non-significant correlation with cooking time under high temperature treatment. ...
Article
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High temperature during the reproductive stage limits the growth and development of lentil (Lens culinaris Medikus). The reproductive and seed filling periods are the most sensitive to heat stress, resulting in limited yield and nutritional quality. Climate change causes frequent incidents of heat stress for global food crop production. This study aimed to assess the impact of high temperature during the reproductive stage of lentil on grain yield, nutritional value, and cooking quality. Thirty-six lentil genotypes were evaluated under controlled conditions for their high temperature response. Genotypic variation was significant (p < 0.001) for all the traits under study. High temperature-induced conditions reduced protein, iron (Fe) and zinc (Zn) concentrations in lentils. Under heat stress conditions, mineral concentrations among lentil genotypes varied from 6.0 to 8.8 mg/100 g for Fe and from 4.9 to 6.6 mg/100 g for Zn. Protein ranged from 21.9 to 24.3 g/100 g. Cooking time was significantly reduced due to high temperature treatment; the range was 3–11min, while under no stress conditions, cooking time variation was from 5 to 14min. Phytic acid variation was 0.5–1.2 g/100 g under no stress conditions, while under heat stress conditions, phytic acid ranged from 0.4 to 1.4 g/100 g. All genotypes had highly significant bioavailable Fe and moderately bioavailable Zn under no stress conditions. Whereas under heat stress conditions, Fe and Zn bioavailability was reduced due to increased phytic acid levels. Our results will greatly benefit the development of biofortified lentil cultivars for global breeding programs to generate promising genotypes with low phytic acid and phytic acid/micronutrient ratio to combat micronutrient malnutrition.
... Many researchers reported negative correlation between seed protein and seed yield and also high heritability (0.84) for protein content [23][24][25]. The differences in both agronomic and seed properties in lentils is the result of genetic and environmental factors [24,[26][27][28][29]. The seed yield is ultimately the result of interaction of genotype with the environment. ...
Article
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Lentil is an important legume crop for human and animal dietary needs due to its high nutritional value. The effect of genotype and growing environment was studied on seed yield (SY), crude protein (CP) and mineral nutrients (macro and micronutrients) of five lentil genotypes grown at four diverse locations for two consecutive years under organic and conventional farming. The location within each year was considered as a separate environment (E). Data were subjected to over environment two-way analysis of variance, while a genotype (G) plus genotype × environment (GGE) biplot analysis was performed. Our results indicated the E as the main source of variation (62.3–99.8%) for SY, CP and macronutrients for both farming systems, while for micronutrients it was either the E or the G × E interaction. Different environments were identified as ideal for the parameters studied: E6 (Larissa/Central Greece/2020) produced the higher CP values (organic: 32.0%, conventional: 27.5%) and showed the highest discriminating ability that was attributed to the lowest precipitation during the crucial period of pod filling. E7 (Thessaloniki/Central Macedonia/2020) and E8 (Orestiada/Thrace/2020) had fertile soils and ample soil moisture and were the most discriminating for high micronutrient content under both farming systems. Location Orestiada showed the highest SY for both organic (1.87–2.28 t ha−1) and conventional farming (1.56–2.89 t ha−1) regardless the year of cultivation and is proposed as an ideal location for lentil cultivation or for breeding for high SY. Genotypes explained a low percentage of the total variability; however, two promising genotypes were identified. Cultivar “Samos” demonstrated a wide adaptation capacity exhibiting stable and high SY under both organic and conventional farming, while the red lentil population “03-24L” showed very high level of seed CP, Fe and Mn contents regardless E or farming system. This genetic material could be further exploited as parental material aiming to develop lentil varieties that could be utilized as “functional” food or consist of a significant feed ingredient.
... The sister lines were selected from the six landraces during IIFC's sesame breeding project in which single plant selection with honeycomb methodology under low plant density (nil-competition) was the core idea. A field evaluation methodology that is advantageous in relation to the elimination of the masking effects of soil heterogeneity, maximization of phenotypic expression and application of high selection intensity was applied [54][55][56]. ...
Article
Full-text available
On-farm genotype screening is at the core of every breeding scheme, but it comes with a high cost and often high degree of uncertainty. Phenomics is a new approach by plant breeders, who use optical sensors for accurate germplasm phenotyping, selection and enhancement of the ge-netic gain. The objectives of this study were to: (1) develop a high-throughput phenotyping workflow to estimate the Normalized Difference Vegetation Index (NDVI) and the Normalized Difference Red Edge index (NDRE) at the plot-level through an active crop canopy sensor; (2) test the ability of spectral reflectance indices (SRIs) to distinguish between sesame genotypes throughout the crop growth period; and (3) identify specific stages in the sesame growth cycle that contribute to phenotyping accuracy and functionality and evaluate the efficiency of SRIs as a selection tool. A diversity panel of 24 sesame genotypes was grown at normal and late planting dates in 2020 and 2021. To determine the SRIs the Crop Circle ACS-430 active crop canopy sensor was used from the beginning of the sesame reproductive stage to the end of the ripening stage. NDVI and NDRE reached about the same high accuracy in genotype phenotyping, even under dense biomass conditions where “saturation” problems were expected. NDVI produced higher broad-sense heritability (max 0.928) and NDRE higher phenotypic and genotypic correlation with the yield (max 0.593 and 0.748, respectively). NDRE had the highest relative efficiency (61%) as an indirect selection index to yield direct selection. Both SRIs had optimal results when the monitoring took place at the end of the reproductive stage and the beginning of the ripening stage. Thus, an active canopy sensor as this study demonstrated can assist breeders to differenti-ate and classify sesame genotypes.
... A different approach from the classical breeding program is the valuable exploitation of the intra-cultivar phenotypic variation through selection at nil competition, as shown in numerous studies on plants such as corn (Zea mays L.) [14], soybean [Glycine max (L.) Merr.] [15], wheat (Triticum aestivum L.) [16], cotton (Gossypium hirsutum L.) [17], snap bean (Phaseolus vulgaris L.) [18] and rice (Oryza sativa L.) [19]. This technique, i.e., the selection at nil competition, is a short-time tool for selection within landraces to breed for high yielding and stable varieties [20][21][22][23][24]. Many projects using molecular techniques have revealed genetic heterogeneity within cultivars [19,25,26]. ...
Article
Full-text available
This study assessed the variations in grain yield (GY) and protein content (PC) within two commercial durum wheat cultivars (Svevo and Maestrale) and evaluated their responses to intra-cultivar selection for both traits. We investigated whether the variations are exploitable and could result in concurrent GY and PC upgrading. The experiments were conducted in the IPBGR, Thessaloniki, Greece (2018–2020). The first year included two identical honeycomb design trials under ultra-low plant density (ULD) where the divergent selection was applied based on single plant yield and protein content. In the second year, progeny evaluation under typical crop density (TCD) for GY and PC occurred in a randomized complete block (RCB) and with three replications for each cultivar selected line. This revealed considerable variation within already improved commercial cultivars. Single-plant selection for GY and PC simultaneously resulted in: (a) one high-yielding line that significantly outperformed the original cultivar Svevo while maintaining high PC, and (b) two high-grain PC lines that outperformed the original cultivar Maestrale significantly while maintaining high GY. ULD allowed efficient selection for GY and PC simultaneously within narrow gene pools by maximizing phenotypic expression and differentiation among individual plants.
... Lentil is a versatile and profitable pulse crop. It is an excellent source of complex carbohydrates, protein, minerals, vitamins, and dietary fiber for humans, and valuable feed and fodder for livestock (Ninou et al., 2019). In India, lentil is a cool-season food legume crop often planted as a rainfed crop during winter. ...
Article
Full-text available
The simultaneous occurrence of high temperature and moisture stress during the reproductive stage of lentil (Lens culinaris Medik) constrains yield potential by disrupting the plant defense system. We studied the detrimental outcomes of heat and moisture stress on rainfed lentils under residual moisture in a field experiment conducted on clay loam soil (Aeric Haplaquept) in eastern India from 2018 to 2019 and from 2019 to 2020 in winter seasons. Lentil was sown on two dates (November and December) to expose the later sowing to higher temperatures and moisture stress. Foliar sprays of boron (0.2% B), zinc (0.5% Zn), and iron (0.5% Fe) were applied individually or in combination at the pre-flowering and pod development stages. High temperatures increased malondialdehyde (MDA) content due to membrane degradation and reduced leaf chlorophyll content, net photosynthetic rate, stomatal conductance, water potential, and yield (kg ha–1). The nutrient treatments affected the growth and physiology of stressed lentil plants. The B+Fe treatment outperformed the other nutrient treatments for both sowing dates, increasing peroxidase (POX) and ascorbate peroxidase (APX) activities, chlorophyll content, net photosynthetic rate, stomatal conductance, relative leaf water content (RLWC), seed filling duration, seed growth rate, and yield per hectare. The B+Fe treatment increased seed yield by 35–38% in late-sown lentils (December). In addition, the micronutrient treatments positively impacted physiological responses under heat and moisture stress with B+Fe and B+Fe+Zn alleviating heat and moisture stress-induced perturbations. Moreover, the exogenous nutrients helped in improving physiochemical attributes, such as chlorophyll content, net photosynthetic rate, stomatal conductance, water potential, seed filling duration, and seed growth rate.
... The results of our study indicated that small progress in grain protein content has been achieved, while selecting for high grain yield, in accordance with Simmonds's (1996) remark. Working with lentil crops, following a 2-year selection cycle for individual plant yield under ultra-low density, Ninou et al. (2019) ensured that the selection of high yielding lines maintained or even improved their seed quality characteristics. ...
Article
Full-text available
Rainfall and temperature are unpredictable factors in Mediterranean environments that result in irregular environmental conditions for crop growth, thus being a critical source of uncertainty for farmers. This study applied divergent single-plant selection for high and low yield within five barley varieties and two Tunisian landraces under semi-arid conditions at an ultra-low density of 1.2 plants/m ² for two consecutive years. Progeny evaluation under dense stands following farmers’ practices was conducted in two semi-arid locations in Tunisia during one cropping season and in one location during a second season, totalling three environments. The results revealed significant genotypic effects for all recorded agronomic and physiological traits. No genotype × environment interaction was shown for biological yield, implying a biomass buffering capacity for selected lines under different environmental conditions. However, genotype × environment interaction was present in terms of grain yield since plasticity for biomass production under drought stress conditions was not translated directly to yield compensation for some of the lines. Nevertheless, several lines selected for high yield were identified to surpass their source material and best checks in each environment, while one line (IH4-4) outperformed consistently by 62.99% on average, in terms of grain yield, the best check across all environments. In addition, improved agronomic performance under drought conditions induced an indirect effect on some grain quality traits. Most of the lines selected for high yield maintained or even improved their grain protein content in comparison to their source material (average increase by 2.33%). On the other hand, most of the lines selected for low yield indicated a poor agronomic performance, further confirming the coherence between selection under ultra-low density and performance under dense stand.
... The methodology applied was either PLS in the case of local populations or BPS. After 2000, lentil breeding is conducted according to the principles of honeycomb selection under nil competition (Iliadis et al. 2003;Vlachostergios et al. 2011Vlachostergios et al. , 2018aVlachostergios and Roupakias 2017;Ninou et al. 2019). Nowadays, national lentil breeding is implemented as a self-funded program by the Institute of Industrial and Forage crops. ...
Chapter
Lentil is an ancient legume crop cultivated thousands of years for its nutritious seeds, its ability to improve soil colonized by nitrogen fixing symbiotic bacteria, and providing income to local farmers at semiarid areas. During the centuries, numerous landraces and traditional varieties have been developed, providing a wealth of genetic material for lentil cultivation and use by local communities worldwide. However, current improved lentil varieties suffer from many biotic and abiotic challenges, and breeding new cultivars should exploit the breadth of genetic potential reserved within the Lens gene pool. Landraces and wild relatives are more tolerant to adverse environmental conditions and can provide valuable genes to develop improved varieties in modern agriculture, adapted to environmental abiotic and biotic stresses, suitable as well for other industrial non-food uses, such as biomass production and use as energy crop. Molecular tools to assist breeding efforts in lentil are less well developed in comparison with other crops, although progress has been made in germplasm characterization using molecular markers. Genomic research is delayed by the large (4.3 GB) lentil genome size, and progress towards the release of the complete lentil genome sequence is expected to accelerate breeding efforts. In this chapter we review current knowledge on lentil domestication and landrace distribution, cultivar improvement and breeding, efforts to characterize abiotic and biotic stress tolerance, the research strategies and major advancements made by modern molecular technologies for identification and utilization of important markers/QTLs in lentil breeding, and future prospects for this important legume crop.
... The methodology applied was either PLS in the case of local populations or BPS. After 2000, lentil breeding is conducted according to the principles of honeycomb selection under nil competition (Iliadis et al. 2003;Vlachostergios et al. 2011Vlachostergios et al. , 2018aVlachostergios and Roupakias 2017;Ninou et al. 2019). Nowadays, national lentil breeding is implemented as a self-funded program by the Institute of Industrial and Forage crops. ...
Chapter
Lentil is an ancient legume crop cultivated thousands of years for its nutritious seeds, its ability to improve soil colonized by nitrogen fixing symbiotic bacteria, and providing income to local farmers at semiarid areas. During the centuries, numerous landraces and traditional varieties have been developed, providing a wealth of genetic material for lentil cultivation and use by local communities worldwide. However, current improved lentil varieties suffer from many biotic and abiotic challenges, and breeding new cultivars should exploit the breadth of genetic potential reserved within the Lens gene pool. Landraces and wild relatives are more tolerant to adverse environmental conditions and can provide valuable genes to develop improved varieties in modern agriculture, adapted to environmental abiotic and biotic stresses, suitable as well for other industrial non-food uses, such as biomass production and use as energy crop. Molecular tools to assist breeding efforts in lentil are less well developed in comparison with other crops, although progress has been made in germplasm characterization using molecular markers. Genomic research is delayed by the large (4.3 GB) lentil genome size, and progress towards the release of the complete lentil genome sequence is expected to accelerate breeding efforts. In this chapter we review current knowledge on lentil domestication and landrace distribution, cultivar improvement and breeding, efforts to characterize abiotic and biotic stress tolerance, the research strategies and major advancements made by modern molecular technologies for identification and utilization of important markers/QTLs in lentil breeding, and future prospects for this important legume crop.
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Lentil (Lens culinaris Medik.) is a self-pollinating diploid (2n=2x=14) species belonging to the Fabaceae family. It is one of the oldest crops known, with 8,000 to 9,000 years of history and it is among the earliest domesticates from the Near East Fertile Crescent. The seeds have high nutritional value. This crop is an interesting substitute to wheat in cereal rotations but its importance is low due to a lack of suitable varieties with local adaptation. Some of the major problems that Argentinian lentil breeders face are the narrow genetic base of the current cultivated germplasm and its low yield potential. A lentil breeding program was initiated in 2004 to develop new varieties with adaptation to prevalent conditions in growing areas of Argentina. Germplasm was obtained from ICARDA (International Center for Agricultural Research in the Dry Areas) and local producers. Conventional breeding methods using hybridization and selection are being carried out to develop improved varieties, broad the genetic base, and isolate superior recombinant inbred lines. Two new varieties have been obtained, one of the macrosperm type (Boyerito FCA) and the other of the microsperm type (Tacuarita FCA) through the application of mass selection in F2 populations from the cross of selected materials. This program complements traditional breeding methods with biotechnological techniques such as transgenesis, use of molecular markers, in vitro embryo culture combined with the SSD method to shorten the breeding time, and digital phenotyping. Key words: Lentil, conventional methodologies, in vitro embryo culture, biotechnology techniques, digital phenotyping.
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In terms of safe food and a healthy food supply, beans (Phaseolus spp.) are a significant source of protein, carbohydrates, vitamins and minerals especially for poor populations throughout the world. They are also rich in unsaturated fatty acids, such as linoleic and oleic acids. From the past to the present, a large number of breeding studies to increase bean yield, especially the common bean (P. vulgaris L.), have resulted in the registration of many modern varieties, although quality and flavor traits in the modern varieties have been mostly ignored. The aim of the present study, therefore, was to compare protein, fat, fatty acid, and some mineral content such as selenium (Se), zinc (Zn) and iron (Fe) of landraces to modern varieties. The landrace LR05 had higher mineral contents, particularly Se and Zn, and protein than the modern varieties. The landrace LR11 had the highest linoleic acid. The landraces are grown by farmers in small holdings for dual uses, such as both dry seed and snap bean production, and are commercialized with a higher cash price. The landraces of the common bean are, not only treasures that need to be guarded for the future, but also important genetic resources that can be used in bean breeding programs. The results of this study suggest that landraces are essential sources of important nutritional components for food security and a healthy food supply.
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Nil-competition (ultra-low plant density) has been asserted to highlight individual genotypes of high yielding potential. This was tested on three lentil (Lens culinaris Medikus) landraces originated from different regions of Greece, germplasm presumably comprising mixtures of homozygous genotypes due to the self-pollinating nature of the crop. Single-plant selection under ultra-low density (interplant distance of 50 or 80 cm) resulted in first- and second-generation sister lines. Progeny testing was conducted in three locations, while the final evaluation at farming density included an additional marginal environment. Wide interplant distance accelerated phenotypic expression of susceptibility to viruses, reflected by high coefficient of variation of single-plant yields. Compared to the mother populations, higher yields combined with reduced virus incidence was observed in the first-generation sister lines, and even higher yields in the second-generation lines partly attributable to further improvement of their sanitary status. Remarkably, at the farming density across five environments, second generation sister lines had mean grain yields by 8, 10 and 20% higher compared to their respective ancestors. Individual sister lines exhibited up to 32% higher yields and stability in ‘agronomic’ terms, i.e. on both the GGE biplot model and regression approach of G×E interaction. In conclusion, the procedure appears an efficient tool that allows the breeder to exploit the natural genetic variability within landraces and develop in short-time pure-line varieties adaptable to a wide range of conditions.
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Lentil is a highly nutritious legume with an ample quantity of carbohydrates and good amount of proteins, minerals, vitamins, phytochemicals and fibres. Although it has been used as staple food since ancient times, its usage has been limited in developed countries, especially due to the lower protein digestibility, presence of anti-nutritional factors, flatulence and poor cooking qualities. Processing of lentils including dehulling and splitting and isolation of major fractions, e.g., proteins and starches are some of the strategies that can be adopted to add value and increase consumption of these legumes. This review paper intends to provide detailed overview of lentil's global production, nutritional composition and processing methods of lentil. Methods of isolation/characterization of lentil protein and starch and their subsequent application in foods are also presented.
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Competition between crop plants, due to resource limitation, is at the root of a considerable yield limitation, a major problem that future agriculture is faced with. Due to inter-plant variation, intra-crop competition causes plant-to-plant interference and unbalanced use of input which decreases possible profit. Acquired intra-crop competition is a priori present in farming due to spatial heterogeneity. Genetically imposed intra-crop competition is due to the plant-to-plant genetic differences, i.e. the intra-species genetic competition in multi-genotypic varieties grown alone, and both the intra- and inter-species genetic competition in intercropping multi-genotypic varieties. In general, high densities accelerate the acquired plant-to-plant variation and intensify the intra-crop competition. Considering environmental diversity, an additional yield gap element is density-reliance. Density-reliant varieties are inefficient in resource use at the single-plant level and present poor results at low densities, accompanied by variation in optimum density particularly in rain-fed agroecosystems. The remedy relies on breeding of varieties that comprise the ‘weak competitor’ ideotype(s) of improved plant yield efficiency in order to mitigate the intra-crop variation and optimise the resource use across variable conditions. To focus on both, selection at nil-competition (widely spaced plants to preclude interference for inputs) is necessary. Selection among spaced plants is further supported by the negative relationship between genotype competitive and yielding ability. The derived density-independent varieties would be capable of taking advantage of the abundance of resources in favourable seasons at low densities that are suitable for dry seasons, approaching the attainable yield across locations and seasons.
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Chickpeas, common beans, dry peas, and lentils are pulse crops that have been a cornerstone of the human diet since the inception of agriculture. However, the displacement of pulses from the diet by low fiber protein alternatives has resulted in a pervasive deficiency referred to as the dietary fiber gap. Using an analytical method American Association of Analytical Chemists (AOAC) 2011.25 that conforms to the Codex Alimentarius Commission consensus definition for dietary fiber, the fiber content of these pulse crops was evaluated in seed types used for commercial production. These pulse crops have 2 to 3 times more fiber per 100 g edible portion than other dietary staples. Moreover, there is marked variation in fiber content among cultivars of the same crop. We conclude that pulse crop consumption should be emphasized in efforts to close the dietary fiber gap. The substantial differences in fiber content among currently available cultivars within a crop can be used to further improve gains in fiber intake without the need to change dietary habits. This provides a rationale for cultivar-based food labeling.
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Landraces are heterogeneous populations and their variability goes through continuous alterations because of physical, genetic, and epigenetic procedures exacerbated by the ongoing climatic changes. Appropriate stewardship of landrace diversity is pivotal to promote its longevity in a manner that is sustainable from the farming perspective. A seed multiplication procedure is presented based on the assumption that in order to improve effectiveness in resource use and increase seed productivity, landraces should comprise genotypes which minimize intra-species competition. These aforementioned genotypes should be of the “weak competitor” ideotype, which are selected so as to alleviate the interplant competition and reach as high as possible crop stand uniformity. Stand uniformity is essential to ensure the same growing conditions for each plant. Reduced intra-crop inequality and equal use of inputs by individual plants will optimize crop performance. Precisely, the “weak competitor” is most often of high yield potential due to a negative association between yielding and competitive ability. Therefore, the suggested procedure involves initial reproduction at nil-competition (widely spaced plants to preclude any plant-to-plant interference for inputs) where “off-type” and low yielding plants are omitted, followed by subsequent multiplication at dense stands. This may represent an effective cultural practice to improve also the landrace health status concerning seed-borne diseases in the absence of certification systems.
Book
Book description: Food legumes are important constituents of human and animal nutrition, supplying high quality proteins crucial for a balanced diet. These crops also play an important role in low-input agricultural production systems by fixing atmospheric nitrogen. Despite systematic and continuous breeding efforts by legume researchers all over the world, substantial genetic gains have not been achieved. These issues require immediate attention, and overall, a paradigm shift is needed in breeding strategies to strengthen our traditional crop improvement programs. To this end, Biology and Breeding of Food Legumes provides extensive information on their history, origin, evolution and botany, as well as breeding objectives and procedures, nutritional improvement, industrial uses, post-harvest technology and recent developments made through biotechnological intervention.
Article
The lentil is among the earliest domesticates from the Near East Fertile Crescent. With seed of high nutritional value and a low water use, the food legume crop – lentils – is well adapted to cereal-based dryland cropping in Mediterranean and subtropical regions. Canada and India are the largest producers. Major production problems addressable through breeding are outlined. To solve these problems, available genetic resources and breeding methodologies for today and tomorrow (biotechnology) are described. Examples of key achievements such as the broadening of the genetic base of the crop in South Asia, the development of plant types suited to a mechanized harvest, and the breeding of lentil with winterhardiness are given. The article concludes by highlighting future challenges in lentil breeding.
Chapter
The lentil is among the earliest domesticates from the Near East Fertile Crescent. With seed of high nutritional value and a low water use, the food legume crop – lentils – is well adapted to cereal-based dryland cropping in Mediterranean and subtropical regions. Canada and India are the largest producers. Major production problems addressable through breeding are outlined. To solve these problems, available genetic resources and breeding methodologies for today and tomorrow (biotechnology) are described. Examples of key achievements such as the broadening of the genetic base of the crop in South Asia, the development of plant types suited to a mechanized harvest, and the breeding of lentil with winterhardiness are given. The article concludes by highlighting future challenges in lentil breeding.