Acta agriculturae Slovenica, 119/1, 1–15, Ljubljana 2023
doi:10.14720/aas.2023.119.1.2970 Original research article / izvirni znanstveni članek
Evaluation of yield and water use efficiency of quinoa under irrigation 
regimes, gamma aminobutyric acid, and vermicompost application
Mehdi JAMSHIDEYNI 1, Mohammad Ali BEHDANI 1, 2, Soheil PARSA 1, Surur KHORAMDEL 3
Received December 22, 2022; accepted March 17, 2023.
Delo je prispelo 22. decembra 2022, sprejeto 17. marca 2023
1 Department of Department of Plant Production and Genetics, Faculty of Agriculture, University of Birjand, Birjand, Iran.
2 Corresponding author, e-mail: mabehdani@birjand.ac.ir
3 Department of Agrotechnology, Faculty of Agriculture, Ferdowsi University of Mashhad, Mashhad, Iran
Evaluation of yield and water use efficiency of quinoa under 
irrigation regimes, gamma aminobutyric acid, and vermi-
compost application
Abstract: The current study was aim to evaluate the in-
teraction effects of gamma aminobutyric acid (GABA) and 
vermicompost on yield and yield components of quinoa 
under different levels of drought stress. For this, two experi-
ments were similarly designed as the factorial-split with four 
replicates for evaluating this hypothesis. Irrigation regimes (50, 
75, and 100 % of plant water requirement (PWR)) as the main 
plot and vermicompost V (0, 5 t ha-1) × gamma aminobutyric 
acid GABA levels (0, 5, 10 mg l-1) as the subplot were designed. 
Severe drought stress had a significant effect on plant height. 
Plant height reduced 31.8 % after using 50 % of PWR compared 
to the control conditions. Although drought stress negatively 
affected the 1000 seed mass and seed yield, GABA foliar ap-
plication alleviated these effects. After using 50 % of PWR, 10 
mg l-1 of GABA increased the seed yield and harvest index up 
to 21.22 and 15.5 %, respectively, compared to the non-foliar 
application. The reduction in PWR from 100 to 50 % led to in-
creasing in P and K concentrations, as well as sugar and proline 
contents. In the same conditions, the use of GABA or V had 
a significant effect on improving these traits. A similar trend 
was also recorded in relation to water use efficiency. Therefore, 
using 10 mg l-1 of GABA and 5 t ha-1 of V can be effective in al-
leviating water stress.
Key words: Chenopodium quinoa; proline; seed yield; 
sugar; drought stress
Abbreviations: V: vermicompost, GABA: gamma amino-
butyric acid, PWR: plant water requirement, WUE: Water use 
efficiency
Ovrednotenje pridelka in učinkovitosti izrabe vode kvinoje 
pri različnih načinih namakanja in dodatku gama aminoma-
slene kisline in komposta deževnikov
Izvleček: Namen raziskave je bil ovrednotiti vzajemne 
učinke gama amino maslene kisline (GABA) in komposta de-
ževnikov (vermikomposta) na pridelek in njegove komponen-
te kvinoje pri različnih ravneh sušnega stresa. V ta namen sta 
bila izvedena dva podobna poskusa kot poskusa z deljenkami 
s štirimi ponovitvami. Načini namakanja (50, 75 in 100 % po-
trebe rastlin po vodi (PWR)) so bili na glavni ploskvi in do-
datki vermikomposta (V; 0, 5 t ha-1) ter gama aminomaslene 
kisline (GABA; (0, 5, 10 mg l-1) na podploskvah. Velik sušni 
stres je imel značilen učinek na višino rastlin. Višina rastlin se je 
zmanjšala za 31,8 % pri 50 % oskrbi rastlin z vodo v primerjavi 
s kontrolo. Čeprav je sušni stres negativno vplival na maso1000 
semen in pridelek semena, je foliaren nanos GABA ublažil te 
učinke. Pri 50 % oskrbi z vodo je dodatek 10 mg l-1 of GABA 
povečal pridelek semena in žeteveni indeks za 21,22 in 15,5 % 
v primerjavi z obravnavanjem brez foliarnega dodatka GABA. 
Zmanjšanje PWR iz 100 na 50 % je vodilo k povečanju kon-
centracij P in K, kot tudi k povečanju vsebnosti sladkorja in 
prolina. Pod enakimi pogoji je imela uporaba GABA ali V zna-
čilen učinek na izboljšanje teh lastnosti. Podoben trend je bil 
zabeležen glede povezave z učinkovitostjo izrabe vode. Zaradi 
vsega naštetega je uporba 10 mg l-1 GABA in 5 t ha-1 V lahko 
učinkovita pri blažitvi sušnega stresa.
Ključne besede: Chenopodium quinoa; prolin; pridelek 
semena; sladkor; sušni stres
Okrajšave: V: vermikompost, GABA: gama amino masle-
na kislina, PWR: zahteva rastline po vodi, WUE: učinkovitost 
izrabe vode
Acta agriculturae Slovenica, 119/1 – 20232
M. JAMSHIDEYNI et al.
1 INTRODUCTION
Quinoa (Chenopodium quinoa Willd.), an annual 
herbaceous plant from the Amaranthaceae family, is 
grown in arid and semi-arid regions of the world (Hino-
josa et al., 2018; Bedoya-Perales et al., 2018), especially 
in Iran (Razzaghi et al., 2012; Ahmadi et al., 2019). The 
crop is well-known for its nutritional values based on the 
carbohydrates, lipids, and proteins nutrients (Gómez‐
Pando et al., 2010). Leucine, phenylalanine, and valine 
(822, 669, 583 mg 100 g-1 of air-dried flour, respectively) 
were determined as the main amino acids of this crop 
(Agza et al., 2018). The grain flour of quinoa has been 
extensively used in the food industry due to its gluten 
free-based values (Wu et al., 2020) and highly nutritive 
quality based on the protein, Ca, Fe, and Zn contents 
(Ballester-Sánchez et al., 2019, Bazile et al., 2016; Präger 
et al., 2018), which can justify the quinoa as a suitable 
product for celiac patients (Peñas et al., 2014).
The appropriate tolerance of quinoa perceiving 
stressful and unfavorable environments were precisely 
reported (Talebnejad and Sepaskhah, 2015). Moreover, 
this plant can be properly considered as a pivotal candi-
date where water shortage is known as a crucial challenge 
(Talebnejad and Sepaskhah, 2015; Bedoya-Peraleset al., 
2018). Ahmadi et al. (2019) reported that quinoa is a 
well-known tolerant crop against the drought stress con-
ditions. However, water shortage and drought stress are 
considered as the serious problems for crop production, 
especially in the east and northeast of Iran (Bannayan et 
al., 2011; Koocheki et al., 2014), the total annual rainfall 
in which is recorded about 187 and 285 mm, respectively, 
indicating water deficiency as the main challenge against 
the successful crop production (Iran meteorological or-
ganization, 2020). Drought stress may significantly af-
fect the plant growth and production by diminishing the 
leaf area (Feng et al., 2013), decelerating the photosyn-
thesis rate (Saeidi et al., 2017; Zhao et al., 2020; Nadali 
et al.,2021), and retarding the enzymatic activity (Xu et 
al., 2015; Askary et al., 2018). Deficit irrigation regimes 
can negatively affect the dry matter production and seed 
yield of quinoa compared to the normal irrigation treat-
ment (Talebnejad and Sepaskhah, 2015). Therefore, it is 
necessary to use the agronomic approaches to increase 
the plant tolerance against the stressful conditions, which 
maximizes the crop yield and water use efficiency (Raz-
zaghi et al., 2012).
Water use efficiency (WUE) is a crucial indicator 
related to evaluating available or applied water that can 
determine an actual plant function, especially in arid 
and semi-arid areas. The concept of WUE indicates how 
much growth and performance changes for each unit of 
water used (Dong et al., 2011). Under dried conditions, 
plants generally use the available water with higher effi-
ciency, although this may lead to a decrease in plant yield 
(Hosseinzadeh et al. 2018). Therefore, when the plant is 
faced with water limitation or suboptimal irrigation, the 
set of operations such as organic matter supply can di-
rectly affect the efficiency of consumed water by stimu-
lating the growth behavior.
Soil fertility can be determined for mitigating the 
stress effects when the plant is affected by the adverse 
environmental conditions. In this regard, the use of ver-
micompost has been known as an appropriate media for 
improving the rhizosphere fertility and increasing the 
crop production (Joshi et al., 2015). Vermicompost can 
significantly promote the soil biological mechanisms 
(Hosseinzadeh et al., 2020) and simultaneously regulate 
the synchrony of minerals uptake during the growing 
seasons due to the balanced nutrient and high organic 
matter contents (Rezaei-Chiyaneh et al., 2020; Xu et al., 
2016). Seyyedi et al. (2016) reported that the expanded 
root growth decreased soil pH, and increased microbial 
activity showed the positive consequences of vermicom-
post application.
Endogenous hormonal modifications are known as 
a plant self-mechanisms for repairing the cellular dam-
age (Dobra et al., 2010), balancing the osmolytes uptake 
(Sarwat and Tuteja, 2017), stabilizing the cell membrane 
(Albacete et al., 2014), and proliferating the plant toler-
ance against the abiotic stresses such as drought condi-
tions (Ni et al., 2013). However, the efficiency of internal 
self-mechanisms may be disturbed by increasing the se-
verity of water shortage. Therefore, the use of exogenous 
phytohormone is possibly useful for mitigating the ad-
verse consequence of water shortage. In this regard, 
γ-aminobutyric acid, a four-carbon non-protein amino 
acid, has been introduced as a crucial plant growth regu-
lator for alleviating the water deficiency during the grow-
ing season (Yong et al., 2017). The use of γ-aminobutyric 
acid (GABA) is proposed for diminishing the side effects 
of water stress by modulating the plant signaling (Ji et al., 
2018), scavenging the reactive oxygen species, and regu-
lating the osmolytes (Vijayakumari and Puthur, 2016), 
which improves the plant growth under water scarcity 
(Rezaei-Chiyaneh et al., 2018); therefore, this study aims 
to evaluate GABA effects on yield and yield components 
of quinoa under different levels of drought stress. Fur-
thermore, the interaction between vermicompost and 
GABA was assessed for finding a suitable strategy in im-
proving water use efficiency.
Acta agriculturae Slovenica, 119/1 – 2023 3
Evaluation of yield and water use efficiency of quinoa ...
2 MATERIALS AND METHODS
2.1 SITE DESCRIPTION
The experiment was conducted in two experimen-
tal stations located in Birjand [Faculty of Agriculture, 
University of Birjand (latitude: 32° 53′N, longitude: 
of 59°13′E, elevation: 1480 m above the sea level)] and 
Mashhad [Faculty of Agriculture, Ferdowsi University 
of Mashhad (latitude: 36◦15′N, longitude: 59◦28′E, el-
evation: 985 m above the sea level)] during the growing 
seasons 2019 (Fig. 1). Both experimental sites were lo-
cated in a semi-arid region, Razavi Khorasan and South 
Khorasan provinces, Iran. Meteorological data during 
the growing season are given in Table 1.
2.2 EXPERIMENTAL DESIGN
Two experiments were similarly designed as the 
factorial-split plot with four replicates. The experimen-
tal factors were included. Irrigation regimes (50, 75, and 
100 % of water requirement) and vermicompost (0, 5 t ha-
1) × GABA (Sigma-Aldrich, A2129; NH2 (CH2)3COOH) 
levels (0, 5, 10 mg l-1) were considered as the main plot 
and subplot, respectively (Fig. 2). The materials related to 
the analyses of vermicompost (produced by Arta Manu-
facturing Group, Iran) included N (1.45 %), P (1.16 %), K 
(1.2 %), and organic matter (18.65 %).
2.3 AGRONOMIC PRACTICES
In both Brijand and Mashhad stations, all agronom-
ic operations were relativity similar. Before preparing the 
seed bed, sampling was done from 0-30 cm depth based 
on the soil of testing location for determining the physi-
cal and chemical properties (Table 2). In the following, 
vermicompost was applied according to experimental 
treatments.
After preparing the experimental soils (deeply 
plowing, twice disking, and leveling), the plots were de-
signed (5 m width and 3 m length). Each block consist-
ed of three main plots with six subplots. The distances 
among the plots and blocks were considered as 0.5 and 
2 m, respectively. Seeds (‘Titicaca’) were sown on 29th 
and 26th July 2019 in Mashhad and Birjand, respectively. 
The final density was determined as 60 plant m-2. Foliar 
spraying of GABA was performed at 2-4 and 10-12 leaf 
stages, respectively.
Based on the weather conditions at both sites, the 
plant water requirement was calculated by using CROP-
WAT software (FAO 1992). The first irrigation was done 
immediately after the seed sowing and the second irriga-
tion was performed three days later. Weed management 
was conducted manually during the growing season. Af-
Month
Minimum temperature 
(oC)
Maximum temperature 
(oC)
Total rainfall 
(mm)
Relative humidity 
(%)
B M B M B M B M
July 21.3 22.4 37.6 37.4 0.0 0.5 13.6 23.6
August 18.8 21.0 36.4 35.5 0.0 0.0 15.1 24.0
September 14.7 15.9 34.0 30.7 0.0 0.0 15.3 26.9
October 10.0 11.7 29.7 26.2 0.0 0.1 18.9 38.4
November 3.9 4.2 20.6 15.8 1.0 0.6 31.9 52.3
Average 13.7 15.0 31.7 29.1 - - 19.0 33.0
Total - - - - 1.0 1.2 - -
30 years 13.0 14.2 30.7 28.9 1.8 5.1 24.6 39.1
Table 1: Weather criteria of experimental sites located at Birjand (B) and Mashhad (M), Iran in 2019
Fig 1: A map graphical presenting the position of the experi-
mental field in Iran. The experiment was conducted in two 
experimental stations located in Birjand and Mashhad
Acta agriculturae Slovenica, 119/1 – 20234
M. JAMSHIDEYNI et al.
ter establishing the plant, the use of drought stress began 
for this purpose. 
2.4 DATA COLLECTION
Six plant plot-1 were randomly selected by consid-
ering the marginal effects (25 cm at the first and end of 
the plots and 2 sides of the rows) at the maturity stage 
(26th and 21st October 2019, in Birjand and Mashhad, re-
spectively). Then, plant height, seed number/plant, seed 
mass/plant and 1000 seed mass were determined. Seed 
and biological yields and harvest index were also meas-
ured at the central part of the plots (11.25 m2). Sugar and 
proline contents were determined based on the studies of 
Dubois et al. (1956) and Bates (1973), respectively, after 
measuring the seed yield. Moreover, P and K concentra-
tions were measured based on the study of Murphy and 
Riley (1962) and Miller (1998).
After determine yield, water use efficiency (WUE) 
was calculated as suggested by Dong et al. (2011). To 
evaluate WUE, total amount of underground water and 
probable run off were to be ignored (Eq. 1).
WUE (kg. m-3) = (grain yield (kg) / total amount of 
water used (m3) x 100
2.5 STATISTICAL ANALYSES
Finally, the measured characteristics were statisti-
cally analyzed. After Bartlett test, data analysis (ANOVA) 
was conducted in split plot factorial design (SPFD) based 
on the randomized complete block design (RCBD) in 
two locations (combined analysis). Means were com-
pared by using LSD test at 5 % probability level. The SAS 
software (Version 9.4) and Excel were used for analyzing 
the data and drawing the figures.
Fig 2: A schematic design of the experiment. Experiments were designed as the factorial-split plot with four replicates. Irrigation 
regimes (50, 75, and 100 % of water requirement) and vermicompost (0 and 5 t ha-1) × GABA levels (0, 5, 10 mg l-1) were consid-
ered as the main plot and subplot, respectively
Experimental sites
% mg/kg
pH EC (dS/m)Clay Silt Sand OC N P K
Birjand 18 12 70 0.16 0.031 11.75 225.3 7.80 4.58
Mashhad 44 32 24 0.48 0.080 6.71 181.5 8.34 1.16
Average 31 22 47 0.32 0.055 9.23 203.4 8.07 2.87
Table 2: The soil quality characteristics of experimental sites located at Birjand and Mashhad, Iran
Acta agriculturae Slovenica, 119/1 – 2023 5
Evaluation of yield and water use efficiency of quinoa ...
Fig 3: Seed and biological yields in quinoa as affected by interaction between vermicompost application and GABA levels
Fig 4: Water use efficiency in quinoa as affected by interaction 
between vermicompost application and GABA levels
role in decreasing the adverse effects of drought stress, 
similar to the GABA effect (Table 3).
Based on the results, the interaction between ir-
rigation and V had a significant effect on seed yield, 
biological yield, and harvest index (Table 5). Thus, the 
amount of seed yield, biological yield, and harvest index 
decreased under the drought stress. However, V reduced 
the negative effects of drought on these indices (Table 5). 
No significant difference was observed between the in-
teraction of 5 t ha-1 of V + 75 % of PWR treatment and 0 
t ha-1 of V + 100 % of PWR treatment as the seed yield. 
Moreover, harvest index in 75 % of PWR + 5 t ha-1 of V 
treatment (21.57 %) was significantly higher than 100 % 
of PWR + 0 t ha-1 of V treatment (20.44 %).
GABA significantly improved the seed yield, biolog-
ical yield, and harvest index, similar to the V use. Under 
severe drought stress, the maximum seed yield (1292.25 
kg ha-1) was obtained by using 10 mg of GABA l-1 (Ta-
ble 6). Similar results were obtained for biological yield 
(6847.38 kg ha-1) and harvest index (18.82 %) (Table 6).  
The maximum seed and biological yields (1724.5 
3 RESULTS
3.1 WEATHER CRITERIA IN BIRJAND AND 
MASHHAD REGIONS
Table 1 indicates the average minimum and maxi-
mum temperatures, as well as total rainfall in Birjand 
and Mashhad regions. During the growing season, the 
average of maximum temperature in Birjand region 
(31.66  oC) was slightly higher than Mashhad region 
(29.12 oC). Similarly, Mashhad region received the higher 
relative humidity (33.03 %) compared to Birjand region 
(19.96 %). Therefore, it seems that quinoa plant was com-
paratively experienced more heat stress under Birjand 
climate.
3.2 YIELD AND YIELD COMPONENT 
As shown in Table 3, the plant height and 1000 
seed mass of quinoa were affected by irrigation regimes, 
GABA, and vermicompost (V) under Birjand and Mash-
had conditions. Irrigation treatments had the significant 
effects on plant height and 1000 seed mass. High level of 
drought stress had a significant negative effect on plant 
height. Thus, the plant height was reduced to 31.8 % after 
using 50 % of PWR compared to the control (Table 3). 
Reduction was observed in Birjand than Mashhad region 
(Table 3). Further, severe drought stress significantly de-
creased 1000 seed mass in Birjand and Mashhad regions. 
Although drought stress caused the negative effects on 
plant height and 1000 seed mass, GABA foliar appli-
cation had the positive effect in reducing these effects. 
Therefore, the use of 5 and 10 mg l-1 GABA increased the 
plant height for 6.8 and 10.5 %, respectively, compared to 
the non-foliar application (Table 3). V played an effective 
Acta agriculturae Slovenica, 119/1 – 20236
M. JAMSHIDEYNI et al.
Treatments
Plant height 
(cm)
1000 seed 
mass (g)
Seed yield 
(kg ha-1)
Biological yield 
(kg ha-1)
Harvest index 
(%)
Location
Birjand 89.81 (12) 1.74 1371.4 7225.2 18.85
Mashhad 104.60 1.82 1554.8 74133.1 20.83
LSD (0.05) 7.179 0.052 251.30 297.49 2.939
Irrigation regimes (%)
50 76.21 1.60 1190.7 6635.6 17.81
75 103.58 1.80 1508.1 7162.8 20.69
100 111.81 1.93 1690.4 8159.1 21.01
LSD (0.05) 5.254 0.071 33.95 120.35 0.355
Vermicompost application (t ha-1)
0 90.60 1.733 1364.4 7068.1 19.11
5 103.83 1.83 1561.7 7570.3 20.57
LSD (0.05) 2.629 0.031 22.03 53.51 0.356
GABA levels (mg l-1)
0 (control) 91.90 1.75 1394.5 7171.4 19.22
5 98.13 1.77 1466.5 7326.8 19.90
10 101.58 1.82 1528.2 7459.3 20.40
LSD (0.05) 3.220 0.038 26.98 66.53 0.437
Average 97.20 1.78 1463.06 7319.15 19.84
S.O.V
Location (L) ** ** NS NS **
R (L) - - - - -
Irrigation regimes (I) ** ** - - -
L × I NS NS ** ** **
R × I (L) - - - - -
GABA levels (G) ** ** ** ** **
Vermicompost application (V) ** ** ** ** **
G × V NS * ** ** **
I × G NS * * ** NS
I × V * NS ** ** **
I × G × V * NS ** ** **
L × G NS NS ** ** **
L × V NS NS NS NS NS
L × G × V NS NS NS NS NS
L × I × G NS NS * * *
L × I × V NS NS NS NS NS
L × I × G × V NS NS NS NS NS
Bartlett’s test for homogeneity 0.51 0.11 0.35 0.84 0.40
Table 3: Plant height, 1000 seed mass, of seed yield, biological yield, and harvest index of quinoa as affected by irrigation regimes, 
vermicompost application, and GABA levels in Birjand and Mashhad, Iran
**: significant at p ≤ 0.01; *: significant at 0.01 < p ≤ 0.05; NS: non-significant (0.05 < p)
Acta agriculturae Slovenica, 119/1 – 2023 7
Evaluation of yield and water use efficiency of quinoa ...
Treatments
Sugar 
(mg 100 g-1)
Proline 
(µmol g-1 FM)
P concentration 
(mg g-1)
K concentration 
(mg g-1)
Water use ef-
ficiency 
(kg m-3)
Location
Birjand 124.75 15.47 0.98 1.58 0.2717
Mashhad 114.34 14.77 0.99 1.59 0.3121
LSD (0.05) 6.860 2.160 0.049 0.110 0.0061
Irrigation regimes (%)
50 125.48 18.36 1.14 1.70 0.3421
75 118.95 15.61 1.03 1.58 0.2894
100 114.20 11.38 0.80 1.47 0.2442
LSD (0.05) 3.580 1.087 0.063 0.089 0.0074
Vermicompost application (t ha-1)
0 118.57 13.59 0.93 1.51 0.2696
5 120.52 16.65 1.05 1.66 0.3142
LSD (0.05) 1.146 0.0347 0.032 0.034 0.0061
GABA levels (mg l-1)
0 (control) 114.84 13.99 0.94 1.53 0.2750
5 119.29 15.10 0.99 1.57 0.2933
10 124.50 16.26 1.04 1.65 0.3037
LSD (0.05) 1.404 0.425 0.039 0.041 0.0074
Average 14.70 15.12 0.99 1.58 0.2919
S.O.V
Location (L) ** ** NS NS **
R (L) - - - - -
Irrigation regimes (I) ** ** ** ** **
L × I NS ** NS NS *
R × I (L) - - - - -
GABA levels (G) ** ** ** ** **
Vermicompost application (V) ** ** ** ** **
G × V NS NS * NS **
I × G ** * * * **
I × V * ** * * **
I × G × V NS NS NS NS **
L × G * NS * * NS
L × V * NS NS NS NS
L × G × V NS NS NS NS NS
L × I × G NS NS NS NS NS
L × I × V NS NS NS NS **
L × I × G × V NS NS NS NS NS
Bartlett’s test for homogeneity 0.52 0.07 0.07 0.21 0.51
Table 4: Some quantity and quality traits of quinoa as affected by irrigation regimes, vermicompost application, and GABA levels 
in Birjand and Mashhad, Iran
**: significant at p ≤ 0.01; *: significant at 0.01 < p ≤ 0.05; NS: non-significant (0.05 < p)
Acta agriculturae Slovenica, 119/1 – 20238
M. JAMSHIDEYNI et al.
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D
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Ta
bl
e 
5:
 P
la
nt
 h
ei
gh
t, 
se
ed
 y
ie
ld
, b
io
lo
gi
ca
l y
ie
ld
, h
ar
ve
st
 in
de
x,
 su
ga
r, 
w
at
er
 u
se
 e
ffi
ci
en
cy
 (W
U
E)
, s
ug
ar
 co
nt
en
t, 
pr
ol
in
e, 
P 
an
d 
K
 co
nc
en
tr
at
io
n 
of
 q
ui
no
a 
as
 a
ffe
ct
ed
 b
y 
in
te
ra
c-
tio
n 
be
tw
ee
n 
irr
ig
at
io
n 
re
gi
m
es
 a
nd
 v
er
m
ic
om
po
st
 ap
pl
ic
at
io
n
Ir
rig
at
io
n 
re
gi
m
es
 (%
)
G
A
BA
 le
ve
ls 
(m
g 
l-1
)
Su
ga
r 
(m
g 
10
0 
g-
1 )
Pr
ol
in
e 
(µ
m
ol
 g
-1
 F
M
)
P (m
g 
g-
1 )
K
 
(m
g 
g-
1 )
W
at
er
 u
se
 
effi
ci
en
cy
 (k
g 
m
-3
)
Se
ed
 y
ie
ld
 
(k
g 
ha
-1
)
Bi
ol
og
ic
al
 y
ie
ld
 
(k
g 
ha
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)
H
ar
ve
st
 
in
de
x 
(%
)
50
0
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8.
63
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9
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7
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1
Ta
bl
e 
6:
 S
ug
ar
 co
nt
en
t, 
pr
ol
in
e, 
P 
an
d 
K
 co
nc
en
tr
at
io
n,
 w
at
er
 u
se
 e
ffi
ci
en
cy
 (W
U
E)
, s
ee
d 
yi
el
d,
 b
io
lo
gi
ca
l y
ie
ld
, a
nd
 h
ar
ve
st
 in
de
x 
of
 q
ui
no
a 
as
 a
ffe
ct
ed
 b
y 
in
te
ra
ct
io
n 
be
tw
ee
n 
irr
ig
at
io
n 
re
gi
m
e 
an
d 
G
A
BA
 le
ve
ls
Acta agriculturae Slovenica, 119/1 – 2023 9
Evaluation of yield and water use efficiency of quinoa ...
and 7794.6 kg ha-1) belonged to 5 t ha-1 of V + 10 mg l-1 
GABA treatment by considering the interaction between 
V and GABA (Fig. 3). Moreover, based on the three-way 
interactions, the highest amount of seed yield, biological 
yield, and harvest index at 50 % of PWR was observed 
in 5 t ha-1 of V + 10 mg l-1 of GABA treatment (Table 7).
In both Mashhad and Brjand regions, the lowest 
seed and biological yield, as well as harvest index, were 
observed in the control treatment; while the highest val-
ues were obtained when 5 t ha-1 V was applied + 10 mg l-1 
GABA. For instance, under Mashhad region, its co-appli-
cation compared to non-application caused an increase 
in seed yield by 28.1 % (Table 8).
3.3 QUALITY TRAITS IN QUINOA SEEDS
Sugar and proline contents, as well as P and K con-
centrations were significantly affected by irrigation re-
gimes, V, and GABA levels (Table 4). The amounts of 
sugar and proline contents increased by reducing PWR 
from 100 to 50 %. In other words, under the high level 
of drought stress, these parameters increased in quinoa 
seeds, which were consistent with results of P and K con-
centrations (Table 4).
The use of V improved the amounts of sugar and 
proline contents as well as P and K concentrations, re-
gardless of drought stress levels. Under the use of 50 % of 
PWR, sugar and proline contents increased in the pres-
ence of 5 t ha-1 of V (Table 5). Similar results were ob-
served after increasing the amount of GABA. In fact, 10 
mg l-1 of GABA led to the best results when 50 % of PWR 
was treated (Table 6). This trend was observed in both 
Mashhad and Birjand regions (Table 9).
3.4 WATER USE EFFICIENCY
According to the results presented in Table 4, the 
interaction of irrigation and V had a significant effect on 
WUE. Irrespective of V application, reducing irrigation 
amount led to an increase in WUE. For example, 50 % 
of PWR compared to 75 % and 100 % of PWR improved 
WUE by 7.3 and 27.8 %, respectively (Table 5).
Similar to the effect of irrigation regime and V, the 
interaction of irrigation and GABA on water use ef-
Irrigation 
regimes 
(%)
GABA 
levels 
(mg l-1)
Vermicompost 
application 
(t ha-1)
Plant 
height (cm)
Seed yield 
(kg ha-1)
Biological 
yield (kg ha-1)
Harvest 
index (%)
Water use 
efficiency (kg m-3)
50 0 
(control)
0 61.76 ± 24.24 836.5 ± 154.55 5916.6 ± 388.5 14.11 ± 2.25 0.239 ± 0.046
5 73.50 ± 10.72 1295.7 ± 162.74 6894.2 ± 198.3 18.77 ± 2.02 0.318 ± 0.053
5 0 75.50 ± 8.83 1101.2 ± 172.14 6439.6 ± 211.1 17.05 ± 2.26 0.343 ± 0.050
5 76.88 ± 8.68 1326.3 ± 148.67 6868.6 ± 229.7 19.29 ± 1.82 0.371 ± 0.049
10 0 82.38 ± 9.68 1191.9 ± 168.55 6572.5 ± 221.9 18.11 ± 2.32 0.381 ± 0.043
5 87.25 ± 10.61 1392.6 ± 225.7 7122.3 ± 311.2 19.53 ± 2.94 0.401 ± 0.066
75 0 0 91.25 ± 8.86 1409.3 ± 161.44 6994.2 ± 244.9 20.12 ± 1.88 0.271 ± 0.033
5 97.00  ±  7.86 1527.8 ± 169.66 7222.8 ± 180 21.12 ± 1.94 0.275 ± 0.033
5 0 100.00  ±  7.78 1439.7 ± 169.75 7045.7 ± 213.2 20.40 ± 2.01 0.285 ± 0.033
5 104.63 ± 8.35 1551.7 ± 161.17 7271.4 ± 122.5 21.32 ± 1.92 0.293 ± 0.035
10 0 112.50  ± 11.12 1486.2 ± 165.71 7117.4 ± 161.1 20.85 ± 1.98 0.299 ± 0.031
5 116.13 ± 10.02 1633.8 ± 202.35 7325.3 ± 381.1 22.27 ± 2.22 0.314 ± 0.041
100 0 0 101.50 ± 8.64 1574.1 ± 188.58 7663.2 ± 168.8 20.52 ± 2.15 0.228 ± 0.029
5 105.38 ± 9.24 1723.6 ± 205.52 8337.2 ± 239.0 20.66 ± 2.25 0.233 ± 0.027
5 0 109.25 ± 9.92 1607.0 ± 179.64 7858.6 ± 196.0 20.43 ± 2.01 0.236 ± 0.027
5 115.38 ± 9.04 1773.2 ± 203.03 8477.1 ± 269.2 20.91 ± 2.24 0.249 ± 0.031
10 0 118.00 ± 8.4 1634.1 ± 180.28 8005.0 ± 320.7 20.39 ± 1.81 0.255 ± 0.030
5 121.38 ± 22.98 1830.6 ± 176.67 8613.2 ± 287.7 21.26 ± 2.00 0.265 ± 0.026
LSD (0.05) 8.050 64.80 152.33 1.062 0.0182
Table 7: Seed yield, biological yield and harvest index of quinoa as affected by interaction between irrigation regimes, GABA 
levels, and vermicompost application
Acta agriculturae Slovenica, 119/1 – 202310
M. JAMSHIDEYNI et al.
ficiency was significant (Table 6). By increasing GABA 
consumption in both levels of V, WUE significantly in-
creased. As an example, 10 mg l-1 of GABA under 50 % 
of PWR compared to no application of GABA led to an 
increase in WUE to 21.9 %. Under 75 and 100 % of PWR, 
a positive effect was also observed in terms of GABA ap-
plication.
By increasing the rate of GABA under both V levels, 
WUE was observed to be significantly increased, so that 
the lowest value was observed in the control treatment 
(no V + no GABA foliar spraying) and the highest value 
(0.33 kg m-3) was observed at 5 t ha-1 of V + 10 mg l-1 of 
GABA (Fig. 4). The results also showed that under severe 
stress (50 % water requirement), the quinoa plant grown 
in Mashhad had a higher WUE than Birjand (Table 10).
4 DISCUSSION
Drought stress is an adverse factor associated with 
the reduced seed yield in crops, which can be imposed 
by unbalanced irrigation (Yuan et al., 2019). Decreased 
plant growth, 1000-seed mass, and harvest index are the 
most important consequences of water stress which were 
Location
GABA 
levels (mg l-1)
Vermicompost 
application (t ha-1)
Seed yield 
(kg ha-1)
Biological 
yield (kg ha-1)
Harvest 
index (%)
Birjand 0 (control) 0 1200.1 ± 310.93 6800.3 ± 747.8 17.41 ± 3.07
5 1295.8 ± 262.05 7043.0 ± 685.8 18.26 ± 2.44
5 0 1347.2 ± 236.42 7117.9 ± 647.1 18.84 ± 2.18
5 1417.4 ± 220.38 7379.9 ± 671.7 19.15 ± 1.95
10 0 1454.3 ± 219.7 7431.2 ± 686.2 19.52 ± 1.9
5 1513.4 ± 255.94 7579.2 ± 791.4 19.92 ± 2.35
Mashhad 0 0 1346.5 ± 405.1 6915.7 ± 845.9 19.10 ± 4.02
5 1469.4 ± 263.1 7186.2 ± 581 20.33 ± 2.37
5 0 1527.6 ± 237.2 7345.3 ± 653.3 20.73 ± 2.1
5 1613.9 ± 243.27 7589.6 ± 662.4 21.21 ± 2.09
10 0 1646.5 ± 247.1 7646.9 ± 782.2 21.49 ± 1.9
5 1724.5 ± 241.94 7794.6 ± 711.4 22.12 ± 2.41
LSD (0.05) 52.91 124.38 0.867
Table 8: Seed yield, biological yield and harvest index of quinoa as affected by interaction between location, irrigation regimes, 
and GABA levels
Location GABA levels (mg l-1) Sugar (mg 100 g-1) P (mg g-1) K (mg g-1)
Birjand 0 (control) 119.98 ± 6.02 0.94 ± 0.16 1.53 ± 0.17
5 124.72 ± 6.25 0.99 ± 0.16 1.57 ± 0.15
10 129.54 ± 8.16 1.03 ± 0.17 1.63 ± 0.16
Birjand 0 109.70 ± 6.54 0.94 ± 0.23 1.52 ± 0.21
5 113.85 ± 6.66 0.99 ± 0.18 1.57 ± 0.15
10 119.47 ± 9.16 1.06 ± 0.19 1.68 ± 0.17
LSD (0.05) 1.828 0.053 0.057
Table 9: Sugar content, P and K concentration of quinoa as affected by interaction between location and GABA levels
Location
Irrigation 
regimes (%)
Water use 
efficiency (kg m-3)
Birjand 50 0.316 ± 0.064
75 0.271 ± 0.029
100 0.228 ± 0.024
Mashhad 50 0.368 ± 0.074
75 0.308 ± 0.032
100 0.260 ± 0.027
LSD (0.05) 0.0105
Table 10: Water use efficiency of quinoa as affected by interac-
tion between location and irrigation regimes
Acta agriculturae Slovenica, 119/1 – 2023 11
Evaluation of yield and water use efficiency of quinoa ...
observed in the plants such as barley and canola (Dreccer 
et al., 2018).
Heat stress, along with drought stress, is another 
disorder in agriculture leading to the reduced crop 
yields, especially in arid and semi-arid regions (Plazas et 
al., 2019). This hypothesis can justify the higher seed and 
biological yields in Mashhad compared to Birjand. Based 
on the results of Table 1, quinoa experienced the higher 
heat stress under Birjand climate compared to Mash-
had climate. Furthermore, the low soil organic matter in 
Birjand compared to the Mashhad region (Table 2) may 
cause the soil conditions to be less favorable for quinoa 
growth. 
Water deficiency led to a decrease in yield indices 
and an increase in seed quality traits. Therefore, it seems 
that accumulated compounds such as proline, protein, 
and sugar acted as a compromising mechanism for im-
proving the adaptation of quinoa to drought conditions 
under severe stress. In this regard, Ebrahimian et al. 
(2019) reported an increase in proline concentration un-
der the drought stress and indicated proline can create 
an internal resistance in the plant under water shortage, 
which is consistent with the results of the study of Naeem 
et al. (2018). These researchers considered the effective 
role of proline in improving the growth, regulating the 
water relations along with the antioxidant activity. Ghaf-
fari et al. (2019) indicated the changes in proline concen-
tration under water-deficient conditions and described 
its changes as a reliable index.
The use of V under drought stress can be effective in 
various aspects. The plants which have a higher ability to 
absorb more nutrients such as V from the soil are more 
tolerant against the drought stress (Demir, 2020; Saba et 
al., 2019). Based on the results of the study of Lim et al. 
(2015), the positive effects of V on increasing the seed 
yield of crops are attributed to the improved soil criteria 
including physical, chemical and biological parameters. 
Aboelsoud and Ahmed (2020) indicated that V increased 
the plant height and 1000-seed mass along with seed yield 
in wheat. Based on the result of this study, V may prolif-
erate the adaptive responses against the drought stress by 
inducing the cellular resistance and increasing the water 
uptake. Furthermore, V caused a significant increase in 
tomato yield under drought stress conditions (Chanda et 
al., 2011). Roberts et al. (2007) reported that V increased 
the number of seeds per plant, plant height, 1000-seed 
mass, and the seed yield in wheat. Therefore, it seems that 
V improved the quinoa seed yield by transition the more 
organic matter to seeds compared to the morphological 
and physiological indices. Li et al. (2019) demonstrated 
that the use of GABA improved the plant tolerance and 
adaptability to stressful conditions by increasing the con-
tent of amino acids. Based on the results of this study, the 
effective role of GABA in rising the values of the men-
tioned parameters is mainly related to the high adapta-
tion of morphological stages to drought stress, especially 
during the pollination stage. Therefore, a positive and 
significant relationship exists between GABA foliar and 
V application, which improves the mechanism of GABA 
in the plant by absorbing the nutrients. The use of V and 
GABA can reduce the consequences of drought stress 
when quinoa is exposed to moderate or intense drought 
stress which is unavoidable in semi-arid climates.
As stated before, increasing irrigation rounds led to 
an increase in WUE. Such behavior is probably achieved 
through further development of the root system which 
can improve the plant’s ability to absorb more available 
water. In this regard, Feizabadi et al. (2021) reported an 
increase in physiological growth, root system develop-
ment, and yield of rapeseed as a result of V application.
The results of the triple-interaction effects can ex-
press a precise concept for the simultaneous application 
of GABA and V, whether under stress or non-stress con-
ditions (Table 16). Under severe stress, co-application of 
5 t ha-1 of V + 10 mg l-1 of GABA recorded the highest 
WUE (0.401 kg m-3). As a new result, a synergistic rela-
tionship can be observed between the simultaneous ap-
plication of GABA and V, which is considered useful for 
dealing with drought stress in quinoa plants.
Under both Birjand and Mashhad regions, a higher 
grain yield was observed following GABA and V. Their 
simultaneous application, as mentioned before, is known 
as the innovation of this research to improve plant resist-
ance against drought stress. Interestingly, simultaneous 
application of GABA and V in Birjand region had a low-
er effect on increasing grain yield than Mashhad region. 
Therefore, it can be concluded that the co-application of 
these treatments may require more suitable conditions 
in terms of temperature to record increasing grain yield. 
Therefore, it can be seen as a crucial relationship between 
the supply of nutrients in the plant and the external en-
vironment.
Along with severe drought to quinoa, the applica-
tion of V played an effective role in increasing WUE. 
This increase was also recorded under mild stress (75 % 
of PWR) and no stress (100 % of PWR). In this regard, 
similar results have been reported in terms of V applica-
tion on increasing WUE in wheat (Azimi et al., 2018). 
Therefore, it can be concluded that the effective role of V 
in increasing WUE has been achieved through the fur-
ther development of the root system and aerial parts. In 
fact, it can be stated that there is a positive and signifi-
cant relationship between plant growth traits and the im-
provement of WUE, which can cause an increase in the 
plant’s ability to deal with drought stress. Generally, any 
factor that can be effective in increasing the plant’s ability 
Acta agriculturae Slovenica, 119/1 – 202312
M. JAMSHIDEYNI et al.
for water absorption can improve WUE (Hosseinzadeh 
et al. 2018). As mentioned earlier, GABA spraying can 
play an effective role in improving growth traits and plant 
height. According to Hosseinzadeh et al. (2018), there is 
a positive relationship between increasing plant growth 
ability and WUE. In fact, under constant water consump-
tion, any factor that led to increasing plant performance 
can improve WUE. Therefore, increasing plant yield after 
spraying GABA will be an effective technique for increas-
ing WUE.
According to Eq. (1), there is a positive relationship 
between plant performance and WUE. Considering the 
similar irrigation scheme applied in both regions, the 
higher WUE in Mashhad may be due to increased plant 
growth and grain yield. In other words, more suitable cli-
matic conditions in the Mashhad region probably pro-
vide the possibility of plant performance in a better way, 
which has subsequently caused more water absorption.
In the present study, no significant difference was 
observed between 75 % of PWR + 5 t ha-1 of V treatment 
and 100  % of PWR treatment based on the seed yield. 
V can decrease the negative effect of moderate drought 
stress by reducing the PWR to 75 %. Thus, the balance 
between the use of V and PWR can be effective in de-
creasing the intense of drought stress. 
5 CONCLUSION
Based on the results, the foliar application of GABA 
can improve the plant height and seed yield of quinoa 
if a relative reduction occurred in PWR. 75 % of PWR 
can be justified as a practical treatment. The same results 
were observed for 1000 seed mass, harvest index, sugar, 
protein, P, and K contents. Moreover, the V treatment 
caused a similar response for improved WUE, which is 
consistent with the use of GABA. The co-application of 
V and GABA was proposed as a new approach in qui-
noa cultivation, which can be particularly interesting in 
semi-arid areas so that farmers can implement it under 
water-limited conditions. Therefore, the use of 10 mg l-1 
of GABA simultaneous with 5 t ha-1 of V can be recom-
mended when the quinoa plant is faced with moderate 
or severe drought stress. However, further research is yet 
needed regarding integrated application of GABA with 
other bio-stimulant inputs in quinoa production. 
6 AUTHORS’ CONTRIBUTION STATE-
MENT
M.A. Behdani and S. Parsa proposed the experi-
mental idea and underwent the all stages of the study. 
M. Jamshideyni conduced the experiment and did the 
sampling data. S. Khoramdel managed data analysis and 
supervised the experiment. Finally, the manuscript was 
proofed by all the authors.
7 DECLARATION OF COMPETING INTER-
EST
The authors declare that they have no known com-
peting financial interests or personal relationships that 
could have appeared to influence the work reported in 
this paper.
8 FUNDING AND/OR CONFLICTS OF IN-
TERESTS/COMPETING INTERESTS
We wish to confirm that there are no known con-
flicts of interest associated with this publication and there 
has been no significant financial support for this work 
that could have influenced its outcome.
9 ACKNOWLEDGEMENTS
This research has been financially supported by 
University of Birjand (Grant Number: 21833/1398). The 
authors gratefully acknowledge the research council of 
University of Birjand.
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