Seed germination and physiological responses of quinoa to selenium priming under drought stress

Gholami, Shocofeh; Dehaghi, Majid Amini; Rezazadeh, Alireza; Naji, Amir Mohammad

doi:10.1590/1678-4499.20210183

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Article • Bragantia 81 • 2022 • https://doi.org/10.1590/1678-4499.20210183 copy

Seed germination and physiological responses of quinoa to selenium priming under drought stress

Authorship SCIMAGO INSTITUTIONS RANKINGS

ABSTRACT

The early stages of quinoa germination are sensitive to drought stress. For this purpose, a study entitled the effect of selenium in different concentrations on germination characteristics and some antioxidant enzymes of quinoa under drought stress conditions with polyethylene glycol (PEG 6000) was investigated. The first experimental factor was seed priming with selenium (from two sources: sodium selenate and selenium nanoparticles: SeNPs ? 33.4 nm) at 0.5, 1.5, 3, 4.5, 6 mg·L^?1 concentrations, besides, no priming treatment was used as control. The second factor was drought stress with PEG 6000 in concentrations 0, –0.4, –0.8, and –1.2 MPa. Drought stress with accumulation of reactive oxygen species (ROS) had a negative effect on most of the measured traits. In seeds that were primed with appropriate selenium concentrations, germination parameters and antioxidant enzyme activity as well as proline and protein content increased compared to the control treatment. Under conditions of severe stress (–1.2 MPa), the highest activity of catalase (CAT), superoxide dismutase (SOD), and ascorbate peroxidase (APX) enzymes was observed in prime with selenium nanoparticles at concentrations of 4.5, 6.0 and 4.5 mg·L^?1, respectively. Concentrations higher than 3 mg·L^?1 of selenium nanoparticles and concentrations of 3 mg·L^?1 sodium selenate had the highest accumulation of photosynthetic pigments under control (stress-free) conditions. The present study shows that selenium priming can reduce the harmful effects of drought stress on quinoa by altering germination properties and biochemical properties.

Key words
antioxidants; Chenopodium quinoa Willd.; photosynthetic efficiency; ROS

SOV	df	Germination percentage	Germination rate	Seedling length	Chlorophyll a content	Chlorophyll b content	Total chlorophyll content	Carotenoid content
Drought stress (D)	3	1752.79^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.454^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	51.841^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.7156^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.4553^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	2.3266^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	2.1716^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Selenium (Se)	1	110.50^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.3861^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.3616^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.14231^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2548^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.7880^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2793^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Selenium concentration (C)	5	911.100^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	14.217^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	14.217^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.14215^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1810^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.60000^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.4448^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
D × Se	3	0.7936^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.447^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.447^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0209^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0175^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.07734^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0586^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
D × C	15	15.052^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.6417^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.6417^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0167^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0157^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.04971^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0179^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Se × C	5	17.322^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1904^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1904^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1073^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1752^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.5524^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2889^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
D × Se × C	15	4.800^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.3351^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.3351^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0076^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0159^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0407^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0261^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Error		2.7301	0.08597	0.0859	0.0010	0.0008	0.0017	0.0036
CV (%)		2.075	0.145	4.366	4.366	4.012	3.988	5.278

SOV	df	Proline content	Protein content	CAT activity	APX activity	SOD activity
Drought stress (D)	3	19.7700^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	32.014^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	17.783^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	22.206^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	24.461^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Selenium (Se)	1	0.9958^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.4209^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	3.5022^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.2481^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	12.270^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Selenium concentration (C)	5	1.1640^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	2.5178^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	3.5091^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	4.2243^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	10.183^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
D × Se	3	0.0245^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0747^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1353^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1239^ns ns represents nonsignificant difference at the 0.05 level of probability.	0.0487 ^ns ns represents nonsignificant difference at the 0.05 level of probability.
D × C	15	0.0693^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0948^* * and ** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.3185^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2495^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.1254*
Se × C	5	1.1094^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	2.0748^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	1.2853^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.8268^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	2.5091^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
D × Se× C	15	0.0585^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0729^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2155^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.0422^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.	0.2090^ * and **** indicate significant differences at p ≤ 0.05 and 0.01, respectively.
Error		0.1776	0.0831	0.0631	0.0690	0.0612
CV (%)		4.799	5.446	4.693	5.961	2.586

Drought stress with PEG
	Chlorophyll a content (mg·g^–1 FW)				Chlorophyll b content (mg·g^–1 FW)
	0 MPa	–0.4 MPa	–0.8 MPa	–1.2 MPa	0 MPa	–0.4 MPa	–0.8 MPa
Control (nonpriming)	0.723 h	0.676 jk	0.589 n	0.389 u	0.522 g	0.402 j	0.274 mn
Nano selenium (mg·L^–1)
0.5	0.755 fg	0.727 h	0.660 k	0.423 s	0.532 g	0.474 i	0.312 l
1.5	0.834 d	0.793 e	0.673 jk	0.503q	0.733 d	0.688 e	0.379 k
3.0	0.876 bc	0.804 e	0.683 j	0.515 q	0.884 b	0.723 de	0.381 k
4.5	0.902 b	0.849 cd	0.685 j	0.547 op	1.003 a	0.735 d	0.285 mn
6.0	0.965 a	0.868 c	0.551 o	0.388 u	0.732 d	0.345 kl	0.264 n
Sodium selenate (mg·L^–1)
0.5	0.744 g	0.719 hi	0.614 m	0.406 t	0.476 i	0.412 j	0.340 kl
1.5	0.784 f	0.737 gh	0.639 l	0.457 rs	0.645 ef	0.550 gef	0.358 kl
3.0	0.864 c	0.783 f	0.654 kl	0.488 r	0.800 c	0.604 ef	0.291 m
4.5	0.740 g	0.703 i	0.588 n	0.369vu	0.588 f	0.411 j	0.271 mn
6.0	0.734 gh	0.683 j	0.541 op	0.359 v	0.432 ij	0.312 l	0.245 no
	Total chlorophyll content (mg·g^-1 FW)				Carotenoid content (mg·g^-1 FW))
Control (nonpriming)	1.23 f	1.05 h	0.95 i	0.715 lm	1.32 de	1.23 e	1.08 f
Nano selenium (mg·L^–1)
0.5	1.26 f	1.08 h	1.05 h	0.772 kl	1.35 de	1.26 e	1.19 ef
1.5	1.36 de	1.22 f	1.08 h	0.944 i	1.66 c	1.53 cd	1.21 e
3.0	1.66 cd	1.46 d	1.28 f	0.810 k	1.88 b	1.58 cd	1.23 e
4.5	1.94 b	1.78 c	1.48 d	1.019 h	2.33 a	1.75 bc	1.37 de
6.0	2.36 a	2.19 ab	1.30 e	0.692 m	1.66 c	1.38 de	1.05 f
Sodium selenate (mg·L^–1)
0.5	1.15 g	1.04 h	1.03h	0.735 l	1.33 de	1.25 e	1.05 f
1.5	1.30 e	1.20 f	1.05 h	0.872 ijk	1.53 cd	1.39 d	1.22 e
3.0	1.55 d	1.38 e	1.14 g	0.923 ij	1.76 bc	1.44 d	1.32 de
4.5	1.30 e	1.20 f	0.92 ij	0.710 lm	1.56 cd	1.38 de	1.05 f
6.0	1.15 g	1.01 h	0.90 j	0.673 n	1.38 de	1.13 ef	0.91 g
	Protein content (%)				Proline content (μmol·g⁻¹ FW)
Control (nonpriming)	2.21ef	1.90 g	1.25 jk	0.88 lm	0.94 rq	1.04 op	1.92 i
Nano selenium
0.5	2.35 e	2.32 e	1.65 hi	1.00 kl	1.08 op	1.54 l	2.01 h
1.5	2.54 cd	2.32 e	1.76 h	1.10 k	1.23 o	1.69 k	2.09 h
3.0	2.84 b	2.64 c	2.00 f	1.35 j	1.32 n	1.71 jk	2.23 g
4.5	3.23 a	2.83 b	2.29 ef	1.37 j	1.36 n	1.73 jk	2.36 f
6.0	2.63 c	2.37 e	1.78 h	1.00 kl	1.47 m	1.82 j	2.85 c
Sodium selenate
0.5	2.26 ef	2.10 ef	1.34 j	0.61 n	1.00 p	1.39 m	1.95 j
1.5	2.44 d	2.26 ef	1.66 hi	0.98 l	1.11 op	1.51 l	2.09 h
3.0	2.74 b	2.36 e	1.57 i	0.94 l	1.20 o	1.56 l	2.13 gh
4.5	2.24 f	2.02 f	1.14 k	0.72 lm	0.98 q	1.13 op	1.94 i
6.0	1.80 g	1.57 i	1.10 k	0.62 n	0.93 rq	1.04 op	1.67 k

Drought stress with PEG
Selenium forms and concentrations	CAT activity (U·min^–1·mg protein)			SOD activity (U·min^–1·mg protein)			APX activity (U·min^–1·mg protein)
Selenium forms and concentrations	0 MPa	–0.4 MPa	–0.8 MPa	–1.2 MPa	0 MPa	–0.4 MPa	–0.8 MPa	–1.2 MPa	0 MPa	–0.4 MPa
Control (non-priming)	0.755 no	1.19 l	1.90 gh	2.07 g	1.27 j	1.55 hi	2.08 f	2.65 cd	0.266 l	0.476 j
Nano selenium
0.5	0.6543 o	1.49 j	2.35 fg	2.45 f	1.32 ij	1.61 h	2.06 f	2.53 cd	0.985 gh	1.112 fg
1.5	0.766 n	1.54 j	2.65 e	2.87 d	1.55 hi	1.95 fg	2.59 cd	3.30 b	1.002 g	1.265 fg
3.0	0.865 mn	1.72 hj	2.85 d	3.05 cd	1.79 gh	2.33 ef	2.76 d	3.35 b	1.043 g	1.352 ef
4.5	0.9856 m	1.80 h	3.35 b	4.70 a	1.88 g	2.62 cd	3.23 bc	3.81 ab	1.332 ef	1.500 de
6.0	1.0067 l	1.30 k	1.85 gh	2.00 g	2.13 ef	2.96 c	3.24 bc	4.03 a	1.00 g	1.232 fg
Sodium selenate
0.5	0.8564 mn	1.42 k	2.24 fg	2.30 fg	1.28 j	1.55 hi	2.09 f	2.50 cd	0.8778 h	1.071 g
1.5	0.7554 no	1.50 j	2.50 ef	2.85 d	1.57 hi	1.84 g	2.48 e	2.99 c	1.004 g	1.221 fg
3.0	0.8342 mn	1.70 i	2.65 e	3.20 c	1.68 h	1.97 fg	2.49 e	3.30 b	0.654 i	0.952 gh
4.5	0.9678 m	1.38 k	1.97 g	2.00 g	1.56 hi	1.88 g	2.37 ef	2.81 cd	0.4431 j	0.664 i
6.0	0.7332 no	0.93 m	1.55 j	1.54 j	1.11 k	1.47 i	1.69 h	2.10 ef	0.2554 l	0.402 k

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[1] *Corresponding author: amini@shahed.ac.ir