Yield components and reproductive, physiological, and root traits used in early selection for nitrogen use efficiency in corn

The objective of this work was to examine the possibility of using yield components and reproductive, physiological, and root traits in early selection for nitrogen use efficiency (NUE) in corn. Sixty-four inbred lines were evaluated under two nitrogen fertilization levels: ideal and low. The evaluations were performed at three phenological stages: eight fully-expanded leaves, tasseling stage, and physiological maturity. It is possible to select superior lines for NUE, but the yield components did not show differential behavior under the different nitrogen levels evaluated. Root, reproductive, and physiological traits are not promising for early selection of corn lines with high NUE. Likewise, the eight-leaves and tasseling stages were not promising for this purpose, since NUE should be estimated taking grain yield into account. However, indirect selection for NUE can be performed via number of ears or using the selection index considering number and weight of ears.

Index terms:
Zea mays; abiotic stress; GBlup; indirect selection; path analysis; selection index

Authorship

Pedro Patric Pinho Morais

Universidade Federal de Viçosa, Departamento de Biologia Geral, Avenida P.H. Rolfs, s/no, Campus Universitário, CEP 36570-900 Viçosa, MG, Brazil. E-mail: pedro.morais@ufv.br, luciano.andrade@ufv.br, borem@ufv.brUniversidade Federal de ViçosaBrazilViçosa, MG, BrazilUniversidade Federal de Viçosa, Departamento de Biologia Geral, Avenida P.H. Rolfs, s/no, Campus Universitário, CEP 36570-900 Viçosa, MG, Brazil. E-mail: pedro.morais@ufv.br, luciano.andrade@ufv.br, borem@ufv.br

Massaine Bandeira e Sousa

Universidade de São Paulo, Escola Superior de Agricultura Luiz de Queiroz, Departamento de Genética, Avenida Pádua Dias, no 11, Caixa Postal 9, CEP 13418-900 Piracicaba, SP, Brazil. E-mail: massaine@usp.br, giovannigalli@usp.br, roberto.neto@usp.brUniversidade de São PauloBrazilPiracicaba, SP, BrazilUniversidade de São Paulo, Escola Superior de Agricultura Luiz de Queiroz, Departamento de Genética, Avenida Pádua Dias, no 11, Caixa Postal 9, CEP 13418-900 Piracicaba, SP, Brazil. E-mail: massaine@usp.br, giovannigalli@usp.br, roberto.neto@usp.br

Giovanni Galli

Luciano Rogério Braatz e Andrade

Roberto Fritsche-Neto

Aluízio Borém de Oliveira

SCIMAGO INSTITUTIONS RANKINGS

Figures | Tables | Formulas

Figures (3)
Tables (5)
Formulas (2)

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Figure 1.
Estimated means for corn (Zea mays) traits under low (LN) and ideal (IN) nitrogen conditions in the environments in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil. NUEY, nitrogen use efficiency based on yield (A); NE, number of ears (B); EL, ear length (C); ED, ear diameter (D); EW, ear weight (E); NRE, number of rows per ear (F); NKR, number of kernels per row (G); NUEVT, nitrogen use efficiency in the VT stage (H); and AxiRL, axial root length (I).

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Figure 2.
GGE biplot analysis of the genotype x environment interaction for 62 corn (Zea mays) inbred lines evaluated in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil, as to: nitrogen use efficiency based on yield (A); number of ears (B); ear length (C); ear diameter (D); ear weight (E); number of rows per ear (F); number of kernels per row (G); total yield (H); and axial root length, which was only evaluated in Anhembi in 2014/2015 and in Piracicaba in 2014/2015 (I). LN, low nitrogen; and IN, ideal nitrogen.

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Figure 3.
GGE biplot analysis of the genotype x environment x nitrogen level interaction, under low (LN) and ideal (IN) nitrogen conditions, for 62 inbred corn (Zea mays) lines evaluated in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil, as to: nitrogen use efficiency based on yield (A); and ear length (B).

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Table 1.
Wald test, as well as mean estimates and coefficient of variation (CV), for fixed effects of intercept, replicates (Rep), block within replicates (Bl/Rep), nitrogen levels (N), environments (Env), and nitrogen levels x environments (N x Env) on traits related to yield components in tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions.

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Table 2.
Wald test, as well as mean estimates and coefficient of variation (CV), for fixed effects of intercept, replicates (Rep), block within replicates (Bl/Rep), nitrogen levels (N), environments (Env), and nitrogen levels x environments (N x Env) on root traits of tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions.

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Table 3.
Maximum likelihood ratio test (LRT) for random effects of genotypes (Gen), genotypes x nitrogen levels (Gen x N), genotypes x environments (Gen x Env), and genotypes x nitrogen levels x environments (Gen x N x Env) on tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions, as well as residue estimates, heritabilities (h²), and Spearman correlation (r_INxLN) between genotypes.

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Table 4.
Direct and indirect effects of low and ideal nitrogen levels on yield components, according to nitrogen use efficiency based on yield (NUEY), nitrogen use efficiency in the V8 stage (NUEV8), and nitrogen use efficiency in the VT stage (NUEVT), for tropical corn (Zea mays) inbred lines.

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Table 5.
Direct and indirect effects of low and ideal nitrogen levels on root, physiological, and reproductive traits, according to nitrogen use efficiency based on yield (NUEY), nitrogen use efficiency in the V8 stage (NUEV8), and nitrogen use efficiency in the VT stage (NUEVT), for tropical corn (Zea mays) inbred lines.

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Figure 1. Estimated means for corn (Zea mays) traits under low (LN) and ideal (IN) nitrogen conditions in the environments in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil. NUEY, nitrogen use efficiency based on yield (A); NE, number of ears (B); EL, ear length (C); ED, ear diameter (D); EW, ear weight (E); NRE, number of rows per ear (F); NKR, number of kernels per row (G); NUEVT, nitrogen use efficiency in the VT stage (H); and AxiRL, axial root length (I).

Figure 2. GGE biplot analysis of the genotype x environment interaction for 62 corn (Zea mays) inbred lines evaluated in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil, as to: nitrogen use efficiency based on yield (A); number of ears (B); ear length (C); ear diameter (D); ear weight (E); number of rows per ear (F); number of kernels per row (G); total yield (H); and axial root length, which was only evaluated in Anhembi in 2014/2015 and in Piracicaba in 2014/2015 (I). LN, low nitrogen; and IN, ideal nitrogen.

Figure 3. GGE biplot analysis of the genotype x environment x nitrogen level interaction, under low (LN) and ideal (IN) nitrogen conditions, for 62 inbred corn (Zea mays) lines evaluated in the municipalities of Anhembi in 2013/2014, of Anhembi in 2014/2015, and of Piracicaba in 2014/2015, in the state of São Paulo, Brazil, as to: nitrogen use efficiency based on yield (A); and ear length (B).

Table 1. Wald test, as well as mean estimates and coefficient of variation (CV), for fixed effects of intercept, replicates (Rep), block within replicates (Bl/Rep), nitrogen levels (N), environments (Env), and nitrogen levels x environments (N x Env) on traits related to yield components in tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions.

Source of variation	Trait⁽¹⁾
Source of variation	DF	NUEY	NE	EL	ED	EW	NRE	NKR	TY	ASI
Intercept	1	1485.39***	3332.5***	28010.3***	41173.0***	6119.2***	3588.2***	17870.9***	2427.75***	655.64***
Rep	1	12.30***	1.50^ns	7.30**	7.00*	38.30***	0.10^ns	1.90^ns	21.69***	5.82*
Bl/Rep	14	9.46^ns	21.00^ns	14.50^ns	10.00^ns	19.40^ns	9.00^ns	21.70^ns	16.76^ns	23.73*
N	1	539.17***	1.10^ns	1.40^ns	1.00^ns	0.20^ns	8.00**	3.40^ns	0.50^ns	0.11^ns
Env	2	420.17***	52.60***	68.40***	206.00***	165.00***	15.00***	121.90***	908.57***	1.62^ns
N x Env	2	132.08***	8.50*	23.10***	43.00***	11.50***	7.00*	9.00*	2.26^ns	0.64^ns
Mean estimates	LN	0.11	17.69	11.58	3.81	60.46	13.20	20.91	2060.13	1.80
Mean estimates	IN	0.04	18.01	11.68	3.83	60.73	13.42	21.22	2037.47	1.83
CV (%)	LN	56.92	37.62	14.69	11.63	28.87	12.31	17.96	68.07	92.86
CV (%)	IN	60.59	38.05	15.31	13.00	30.95	11.94	18.57	67.39	89.55

⁽¹⁾
NUEY, nitrogen use efficiency based on yield; NE, number of ears; EL, ear length; ED, ear diameter; EW, ear weight; NRE, number of rows per ear; NKR, number of kernels per row; TY, total yield; and ASI, anthesis-silking interval. DF, degrees of freedom. ***, **, and *Significant at 0.1, 1.0, and 5.0% probability, respectively. ^nsNonsignificant.

Table 2. Wald test, as well as mean estimates and coefficient of variation (CV), for fixed effects of intercept, replicates (Rep), block within replicates (Bl/Rep), nitrogen levels (N), environments (Env), and nitrogen levels x environments (N x Env) on root traits of tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions.

Source of variation	Trait⁽¹⁾
Source of variation	DF	LatRL	AxiRL	LatRV	AxiRV	PEPSII	NUEV8	NUEVT
Intercept	1	3501.30***	4240.40***	2859.04***	3165.60***	37178.00***	5077.90***	5852.10***
Rep	1	4.50*	5.20*	4.24*	0.20^ns	7.00**	7.80**	1.50^ns
Bl/Rep	14	17.20^ns	12.40^ns	10.65^ns	12.70^ns	26.00*	14.00^ns	23.30^ns
N	1	5.00*	6.10*	2.83^ns	4.80*	129.00***	1282.10***	1678.50***
Env	1	63.10***	37.80***	61.41***	77.40***	5.00*	305.20***	26.30***
N x Env	1	1.10^ns	3.80*	1.19^ns	2.00^ns	1.00^ns	126.10***	18.00***
Mean estimates	LN	878.28	645.08	0.43	42.43	0.66	0.01	0.07
Mean estimates	IN	821.15	606.69	0.41	39.70	0.74	0.03	0.22
Coefficient of variation (%)	LN	41.37	32.55	43.46	40.33	12.50	35.06	30.74
Coefficient of variation (%)	IN	43.28	34.36	42.21	41.91	9.81	37.41	22.59

⁽¹⁾
LatRL, lateral root length; AxiRL, axial root length; LatRV, lateral root volume; AxiRV, axial root volume; PEPSII, photosynthetic efficiency of photosystem II; NUEV8, nitrogen use efficiency in the V8 stage; and NUEVT, nitrogen use efficiency in the VT stage. DF, degrees of freedom. ***, **, and *Significant at 0.1, 1.0, and 5.0% probability, respectively. ^nsNonsignificant.

Table 3. Maximum likelihood ratio test (LRT) for random effects of genotypes (Gen), genotypes x nitrogen levels (Gen x N), genotypes x environments (Gen x Env), and genotypes x nitrogen levels x environments (Gen x N x Env) on tropical corn (Zea mays) inbred lines under low (LN) and ideal (IN) nitrogen conditions, as well as residue estimates, heritabilities (h²), and Spearman correlation (r_INxLN) between genotypes.

Source of variation		Trait⁽¹⁾
Source of variation		NUEY	NE	EL	ED	EW	NRE	NKR	TY
Gen		16.20***	60.81***	93.57***	74.2***	70.05***	93.92^***	92.93***	38.5***
Gen x N		13.38***	0.00^ns	0.00^ns	0.00^ns	0.65^ns	0.00^ns	0.00^ns	0.00^ns
Gen x Env		18.36***	43.65***	6.01**	44.7***	102.34***	20.27***	24.75***	95.6***
Gen x N x Env		21.41***	0.64^ns	5.41*	0.00^ns	0.00^ns	1.83^ns	0.02^ns	0.00^ns
Residue		4.3x10^-4	16.42	1.08	6.2x10^-2	70.36	0.85	5.07	219929.70
h²	LN	0.70	0.66	0.73	0.68	0.79	0.78	0.76	0.68
h²	IN	0.67	0.71	0.69	0.73	0.69	0.81	0.69	0.65
r_INxLN	-	0.87	0.82	0.86	0.89	0.90	0.89	0.87	0.87
		ASI	LatRL	AxiRL	LatRV	AxiRV	PEPSII	NUEV8	NUEVT
Gen		45.27***	51.92***	23.79***	45.19***	20.34***	0.00^ns	4.89*	5.07*
Gen x N		8.87***	0.00^ns	0.00^ns	0.00^ns	0.00^ns	0.10^ns	0.00^ns	3.93*
Gen x Env		0.05^ns	0.06^ns	2.89*	0.61^ns	0.70^ns	0.21^ns	0.90^ns	0.00^ns
Gen x N x Env		0.00^ns	0.00^ns	0.00^ns	0.10^ns	0.12^ns	0.01^ns	0.00^ns	0.25^ns
Residue		1.51	75971.67	28747.95	0.02	184.08	5.6x10^-3	5.2x10^-5	1.0x10^-3
h²	LN	0.62	0.45	0.36	0.47	0.46	0.11	0.31	0.08
h²	IN	0.69	0.51	0.28	0.49	0.14	0.00	0.12	0.37
r_INxLN	-	0.62	0.81	0.71	0.78	0.71	0.15	0.59	0.36

⁽¹⁾
NUEY, nitrogen use efficiency based on yield; NE, number of ears; EL, ear length; ED, ear diameter; EW, ear weight; NRE, number of rows per ear; NKR, number of kernels per row; TY, total yield; ASI, anthesis-silking interval; LatRL, lateral root length; AxiRL, axial root length; LatRV, lateral root volume; AxiRV, axial root volume; PEPSII, photosynthetic efficiency of photosystem II; NUEV8, nitrogen use efficiency in the V8 stage; and NUEVT, nitrogen use efficiency in the VT stage. ***, **, and *Significant at 0.1, 1.0, and 5.0% probability, respectively. ^nsNonsignificant.

Table 4. Direct and indirect effects of low and ideal nitrogen levels on yield components, according to nitrogen use efficiency based on yield (NUEY), nitrogen use efficiency in the V8 stage (NUEV8), and nitrogen use efficiency in the VT stage (NUEVT), for tropical corn (Zea mays) inbred lines.

Effect⁽¹⁾		Low N			Ideal N
Effect⁽¹⁾		NUEY	NUEV8	NUEVT	NUEY	NUEV8	NUEVT
	Number of ears (NE)
Direct		0.77	0.07	0.10	0.77	0.11	-0.34
Indirect
	EL	0.01	0.01	0.03	0.00	0.00	-0.03
	ED	0.00	-0.01	0.00	0.02	0.05	0.06
	EW	-0.08	0.02	0.02	-0.05	0.00	0.02
	NRE	0.00	0.00	0.02	0.00	-0.01	0.01
	NKR	-0.02	0.02	-0.01	-0.02	-0.02	0.30
	TY	-	0.03	-0.06	-	0.04	0.16
Total		0.68	0.14	0.10	0.72	0.17	0.18
	Ear length (EL)
Direct		0.07	0.05	0.21	-0.02	-0.03	-0.14
Indirect
	NE	0.13	0.01	0.01	0.17	0.02	-0.07
	ED	0.02	0.18	0.11	0.07	0.20	0.22
	EW	0.33	-0.07	-0.06	0.24	0.01	-0.09
	NRE	0.00	0.00	0.01	0.00	0.00	0.01
	NKR	-0.03	0.03	-0.01	-0.02	-0.02	0.31
	TY	-	0.02	-0.05	-	0.02	0.11
Total		0.52	0.22	0.22	0.44	0.20	0.35
	Ear diameter (ED)
Direct		0.07	0.53	0.32	0.13	0.38	0.41
Indirect
	NE	-0.02	0.00	0.00	0.12	0.02	-0.05
	EL	0.02	0.01	0.07	-0.01	-0.02	-0.07
	EW	0.46	-0.10	-0.09	0.32	0.01	-0.11
	NRE	0.00	-0.01	-0.07	0.01	0.05	-0.04
	NKR	-0.01	0.01	-0.01	-0.01	-0.01	0.20
	TY	-	0.02	-0.05	-	0.03	0.13
Total		0.52	0.46	0.17	0.56	0.46	0.47
	Ear weight (EW)
Direct		0.62	-0.14	-0.13	0.54	0.02	-0.19
Indirect		-	-	-	-	-	-
	NE	-0.10	-0.01	-0.01	-0.08	-0.01	0.03
	EL	0.03	0.02	0.11	-0.01	-0.01	-0.06
	ED	0.05	0.39	0.24	0.08	0.23	0.25
	NRE	0.00	0.00	-0.04	0.01	0.04	-0.03
	NKR	-0.02	0.02	-0.01	-0.02	-0.02	0.21
	TY	-	0.02	-0.05	-	0.03	0.12
Total		0.58	0.30	0.11	0.52	0.28	0.33
	Number of rows per ear (NRE)
Direct		0.00	-0.03	-0.20	0.03	0.17	-0.14
Indirect
	NE	-0.08	0.00	-0.01	-0.06	-0.01	0.02
	EL	0.00	0.00	-0.01	0.00	0.00	0.00
	ED	0.03	0.20	0.12	0.04	0.11	0.12
	EW	0.11	-0.03	-0.02	0.14	0.01	-0.05
	NKR	0.00	0.00	0.00	-0.01	-0.01	0.09
	TY	-	0.00	0.00	-	0.01	0.03
Total		0.04	0.14	-0.12	0.14	0.28	0.07
	Number of kernels per row (NKR)
Direct		-0.04	0.05	-0.03	-0.04	-0.04	0.52
Indirect		-	-	-	-	-	-
	NE	0.34	0.03	0.04	0.44	0.06	-0.19
	EL	0.04	0.03	0.13	-0.01	-0.02	-0.08
	ED	0.02	0.13	0.08	0.05	0.15	0.16
	EW	0.25	-0.05	-0.05	0.21	0.01	-0.08
	NRE	0.00	0.00	0.00	0.01	0.03	-0.02
	TY	-	0.02	-0.05	-	0.04	0.15
Total		0.61	0.21	0.12	0.66	0.23	0.46
	Total yield (TY)
Direct		-	0.04	-0.08	-	0.06	0.22
Indirect		-	-	-	-	-	-
	NE	-	0.05	0.07	-	0.08	-0.24
	EL	-	0.03	0.11	-	-0.02	-0.06
	ED	-	0.27	0.17	-	0.22	0.23
	EW	-	-0.08	-0.08	-	0.01	-0.10
	NRE	-	0.00	0.00	-	0.02	-0.02
	NKR	-	0.03	-0.02	-	-0.02	0.36
Total		-	0.33	0.17	-	0.35	0.39
R²		0.93	0.24	0.11	0.87	0.25	0.33
P_ɛ		0.25	0.87	0.95	0.36	0.86	0.81

⁽¹⁾
R2, Coefficient of determination; and Pɛ, residual effect.

Table 5. Direct and indirect effects of low and ideal nitrogen levels on root, physiological, and reproductive traits, according to nitrogen use efficiency based on yield (NUEY), nitrogen use efficiency in the V8 stage (NUEV8), and nitrogen use efficiency in the VT stage (NUEVT), for tropical corn (Zea mays) inbred lines.

Effect⁽¹⁾		Low N			Ideal N
Effect⁽¹⁾		NUEY	NUEV8	NUEVT	NUEY	NUEV8	NUEVT
	Lateral root length (LatRL)
Direct		-0.33	0.18	0.52	-0.56	0.30	0.52
Indirect
	AxiRL	-0.19	0.00	0.10	-0.28	-0.23	0.06
	LatRV	0.39	-0.21	-0.52	0.82	-0.10	-0.59
	AxiRV	0.14	0.34	0.09	0.10	0.36	0.18
	PEPSII	0.00	0.00	0.00	0.01	0.00	0.00
	ASI	0.00	0.00	0.00	0.00	0.00	0.00
Total		0.01	0.31	0.19	0.09	0.33	0.17
Axial root length (AxiRL)
Direct		0.26	0.00	0.14	-0.40	-0.31	0.09
Indirect
	LatRL	-0.24	0.14	0.39	-0.40	0.21	0.37
	LatRV	0.37	-0.20	-0.48	0.76	-0.10	-0.55
	AxiRV	0.20	0.47	0.12	0.14	0.48	0.25
	PEPSII	-0.01	0.00	-0.01	0.02	0.00	0.00
	ASI	-0.01	0.00	-0.01	0.01	-0.02	0.00
Total		0.57	0.41	0.15	0.13	0.26	0.16
Lateral root volume (LatRV)
Direct		0.43	-0.23	-0.56	0.88	-0.12	-0.64
Indirect
	LatRL	-0.30	0.17	0.48	-0.52	0.28	0.48
	AxiRL	-0.22	0.00	0.12	-0.34	-0.27	0.07
	AxiRV	0.16	0.38	0.10	0.12	0.40	0.21
	PEPSII	0.00	0.00	0.00	0.01	0.00	0.00
	ASI	-0.01	0.00	-0.01	0.00	-0.01	0.00
Total		0.06	0.32	0.13	0.15	0.28	0.12
Axial root volume (AxiRV)
Direct		0.28	0.68	0.18	0.20	0.68	0.35
Indirect
	LatRL	-0.16	0.09	0.27	-0.29	0.16	0.27
	AxiRL	-0.18	0.00	0.09	-0.28	-0.22	0.06
	LatRV	0.24	-0.13	-0.32	0.52	-0.07	-0.37
	PEPSII	-0.02	0.00	-0.02	0.03	0.00	0.00
	ASI	0.00	0.00	0.00	0.00	0.00	0.00
Total		0.16	0.64	0.20	0.18	0.55	0.31
Photosynthetic efficiency of photosystem II (PEPSII)
Direct		0.11	-0.01	0.09	0.13	0.00	0.00
Indirect
	LatRL	0.01	0.00	-0.02	-0.06	0.03	0.06
	AxiRL	0.02	0.00	-0.01	-0.07	-0.05	0.01
	LatRV	-0.01	0.00	0.01	0.09	-0.01	-0.06
	AxiRV	-0.06	-0.14	-0.04	0.04	0.14	0.07
	ASI	-0.01	0.00	-0.01	0.00	0.00	0.00
Total		-0.06	-0.15	0.02	0.13	0.11	0.08
Anthesis-silking interval (ASI)
Direct		-0.07	0.00	-0.04	0.02	-0.12	0.00
Indirect
	LatRL	-0.03	0.02	0.05	-0.02	0.01	0.02
	AxiRL	-0.04	0.00	0.02	-0.07	-0.06	0.01
	LatRV	0.06	0.01	-0.08	0.10	-0.01	-0.07
	AxiRV	0.00	0.01	0.00	0.00	-0.01	0.00
	PEPSII	0.02	0.00	0.01	0.00	0.00	0.00
Total		-0.06	-0.02	-0.04	0.03	-0.19	-0.04
R²		0.06	0.42	0.09	0.08	0.39	0.14
P_ɛ		0.96	0.76	0.95	0.95	0.78	0.93

⁽¹⁾
R²,coefficient of determination; and Pɛ, residual effect.

ρ = 1 - \frac{6 \sum d_{i}^{2}}{n (n^{2} - 1)}

{\hat{h}}^{2} = \frac{σ_{g}^{2}}{σ_{f}^{2}}

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Yield components and reproductive, physiological, and root traits used in early selection for nitrogen use efficiency in corn

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[TFN1] ⁽¹⁾
NUEY, nitrogen use efficiency based on yield; NE, number of ears; EL, ear length; ED, ear diameter; EW, ear weight; NRE, number of rows per ear; NKR, number of kernels per row; TY, total yield; and ASI, anthesis-silking interval. DF, degrees of freedom. ***, **, and *Significant at 0.1, 1.0, and 5.0% probability, respectively. ^nsNonsignificant.