Growth and Physiology Vary Little Among North American, an Asian, and a North American x Asian Hybrid Ash in a Common Garden in Ohio, U.S.A

Laurel J. Haavik; Daniel A. Herms

doi:10.48044/jauf.2019.008

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Figure 1.
Precipitation (gray bars) and minimum (dotted line) and maximum (solid line) temperatures during the 2015 growing season in Wooster, Ohio. Arrows indicate dates on which gas exchange physiological measurements of ash trees were taken.
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Figure 2.
(a) Diameter (at breast height), (b) Specific leaf area (SLA), and (c) N per unit leaf area (LA) by ash taxa. Dots represent means of eight trees (seven for Man x black), and error bars are one standard error from the mean. Different letters indicate significant differences according to Tukey’s HSD. Man × black = Manchurian × black ash.
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Figure 3.
Mean (a) CO₂ assimilation rate (A), (b) stomatal conductance (g_s), (c) photosynthetic nitrogen use efficiency (PNUE), and (d) efficiency of photosystem II (ɸPSII) for six ash taxa by sampling date. Key for taxa is located in (b). Dots represent means of eight trees, and error bars are one standard error from the mean. Man x black = Manchurian x black ash.

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Table 1.

Ash (Fraxinus) species, cultivar, and diameter at breast height (dbh ± standard error) of trees at the start of the 2015 growing season in Wooster, Ohio, U.S.A.

Ash species	Cultivar	Sample Mean	size dbh (cm)
Green, F. pennsylvanica	Patmore	8	3.91 ± 0.71
White, F. americana	Autumn Purple	8	1.38 ± 0.07
Black, F. nigra	Fall Gold	8	4.02 ± 0.31
Blue, F. quadrandgulata	seedling origin	8	1.98 ± 0.16
Manchurian × black, F. mandshurica × F. nigra	Northern Treasure	8	1.56 ± 0.16
Manchurian, F. mandshurica	seedling origin	8	1.90 ± 0.10

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Table 2.

Pearson’s correlation coefficients for leaf traits, growth, photosynthesis, stomatal conductance, and fluorescence among all trees on June 22, 2015 (=wet).

Blue	SLA	dia gr	A	PNUE	g_s	C_i	Fv’/Fm’	ΦPSII
N	0.93^***	0.46	−0.45	−0.92^**	−0.24	−0.04	−0.56	−0.69
SLA		0.31	−0.63	−0.95^***	−0.54	−0.28	−0.77^*	−0.66
dia gr			−0.14	−0.48	−0.11	−0.18	0.33	−0.40
A				0.72^*	0.75^*	0.16	0.57	0.77^*
PNUE					0.53	0.20	0.60	0.79^*
g_s						0.74^*	0.60	0.27
C_i							0.30	−0.28
Fv’/Fm’								0.31

White
N	0.99^***	−0.58	−0.57	−0.82^*	0.16	0.46	−0.64	−0.37
SLA		−0.54	−0.61	−0.84^**	0.16	0.51	−0.67	−0.44
dia gr			0.12	0.36	0.07	0.11	0.20	−0.03
A				0.78^*	0.21	−0.60	0.50	0.71^*
PNUE					0.31	−0.30	0.77^*	0.41
g_s						0.63	0.36	−0.22
C_i							−0.06	−0.78^*
Fv’/Fm’								0.04

Green
N	0.99^***	−0.12	−0.55	−0.88^**	−0.57	0.24	−0.05	−0.01
SLA		−0.20	−0.56	−0.85^**	−0.57	0.25	−0.06	0.02
dia gr			−0.07	−0.56	0.19	0.02	−0.47	−0.43
A				0.82^*	0.67	−0.75^*	0.08	0.71^*
PNUE					0.73^*	−0.46	0.02	0.41
g_s						−0.04	0.38	0.41
C_i							0.20	−0.59
Fv’/Fm’								0.35

Black
N	0.98^***	−0.40	0.26	−0.60	0.23	−0.05	0.37	0.31
SLA		−0.31	0.08	−0.73^*	0.06	−0.21	0.21	0.15
dia gr			−0.14	0.29	−0.41	−0.23	−0.14	0.08
A				0.54	0.85^**	0.63	0.93^***	0.80^*
PNUE					0.44	0.50	0.40	0.31
g_s						0.87^**	0.91^**	0.42
C_i							0.74^*	0.27
Fv’/Fm’								0.65

Man × Black
N	0.67	−0.31	−0.28	−0.72	0.01	0.36	0.09	0.05
SLA		−0.09	−0.59	−0.76^*	−0.25	0.43	−0.38	−0.27
dia gr			−0.01	−0.01	−0.15	−0.24	−0.27	−0.45
A				0.85^*	0.77^*	−0.13	0.63	0.81^*
PNUE					0.54	−0.27	0.42	0.60
g_s						0.53	0.91^**	0.85^**
C_i							0.54	0.23
Fv’/Fm’								0.77^*

Manchurian
N	0.83^**	0.74^*	0.44	−0.21	0.23	0.11	0.69	−0.18
SLA		0.43	0.01	−0.57	−0.01	−0.05	0.48	−0.43
dia gr			0.52	0.06	0.15	−0.10	0.52	−0.18
A				0.78^*	0.79^*	0.57	0.53	0.55
PNUE					0.74^*	0.60	0.14	0.71^*
g_s						0.94^***	0.66	0.48
C_i							0.63	0.41
Fv’/Fm’								−0.20

↵*P < 0.05,
↵**P < 0.01,
↵***P < 0.001; N = leaf nitrogen (μmol mol⁻² s⁻¹); SLA = specific leaf area (m²g⁻¹); dia gr = diameter growth in 2015 (cm); A = net assimilation rate (μmol m⁻² s⁻¹); g_s = stomatal conductance (mol m⁻² s⁻¹); PNUE = photosynthetic nitrogen use efficiency (μmol mol⁻² s⁻¹); C_i = intercellular CO₂ (μmol mol⁻¹); Fv’/Fm’ = light-adapted variable fluorescence; ΦPSII = fraction of absorbed photons used by photosystem II.

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Table 3.

Pearson’s correlation coefficients for leaf traits, growth, photosynthesis, stomatal conductance, and fluorescence among all trees on July 27, 2015 (=wet).

Blue	SLA	dia gr	A	PNUE	g_s	C_i	Fv’/Fm’	ΦPSII
N	0.93^***	0.46	0.05	−0.79^*	−0.03	−0.11	−0.48	−0.09
SLA		0.31	−0.14	−0.83^**	−0.35	−0.29	−0.48	0.09
dia gr			−0.19	−0.57	0.16	0.29	−0.28	0.04
A				0.55	0.63	−0.33	0.60	0.91^**
PNUE					0.34	−0.15	0.72^*	0.46
g_s						0.52	0.86^**	0.46
C_i							0.42	−0.45
Fv’/Fm’								0.37

White
N	0.99^***	−0.58	−0.56	−0.84^**	0.43	0.77^*	−0.51	−0.35
SLA		−0.54	−0.54	−0.84^**	0.48	0.80^*	−0.44	−0.32
dia gr			0.44	0.45	0.06	−0.25	0.79^*	0.42
A				0.76^*	0.20	−0.62	0.69	0.83^*
PNUE					−0.21	−0.77^*	0.61	0.67
g_s						0.64	0.19	0.54
C_i							−0.35	−0.23
Fv’/Fm’								0.59

Green
N	0.99^***	−0.12	0.49	−0.97^***	0.42	0.02	0.44	0.42
SLA		−0.20	0.42	−0.95^***	0.37	0.01	0.45	0.45
dia gr			−0.17	0.04	0.36	0.52	−0.58	−0.39
A				−0.40	0.27	−0.43	0.75^*	0.51
PNUE					−0.39	−0.06	−0.37	−0.29
g_s						0.74^*	−0.03	0.32
C_i							−0.62	−0.13
Fv’/Fm’								0.62

Black
N	0.98^***	−0.40	0.19	−0.77^*	0.08	−0.05	0.21	−0.20
SLA		−0.31	0.08	−0.83^*	−0.06	−0.21	0.03	−0.31
dia gr			−0.05	0.33	−0.30	−0.55	0.04	0.31
A				0.38	0.91^**	0.63	0.76^*	0.71^*
PNUE					0.50	0.48	0.28	0.54
g_s						0.88^**	0.68	0.62
C_i							0.56	0.50
Fv’/Fm’								0.82^*

Man × Black
N	0.67	−0.31	0.38	−0.30	0.19	−0.11	0.67	0.61
SLA		−0.09	0.29	−0.16	−0.09	−0.43	0.48	0.69
dia gr			0.54	0.80^*	0.52	0.29	0.27	0.17
A				0.76^*	0.79^*	0.38	0.77^*	0.56
PNUE					0.68	0.44	0.31	0.24
g_s						0.86^**	0.46	−0.03
C_i							0.06	−0.52
Fv’/Fm’								0.72^*

Manchurian
N	0.83^**	0.74^*	0.40	−0.08	−0.10	−0.45	0.44	0.25
SLA		0.43	0.21	−0.22	−0.36	−0.50	0.12	0.18
dia gr			0.41	0.09	0.34	0.14	0.77^*	0.03
A				0.88^**	0.76^*	0.11	0.60	0.75^*
PNUE					0.88^**	0.34	0.42	0.70
g_s						0.65	0.63	0.47
C_i							0.46	−0.16
Fv’/Fm’								0.01

↵*P < 0.05,
↵**P < 0.01,
↵***P < 0.001; N = leaf nitrogen (μmol mol⁻² s⁻¹); SLA = specific leaf area (m²g⁻¹); dia gr = diameter growth in 2015 (cm); A = net assimilation rate (μmol m⁻² s⁻¹); g_s = stomatal conductance (mol m⁻² s⁻¹); PNUE = photosynthetic nitrogen use efficiency (μmol mol⁻² s⁻¹); C_i = intercellular CO₂ (μmol mol⁻¹); Fv’/Fm’ = light-adapted variable fluorescence; ΦPSII = fraction of absorbed photons used by photosystem II.

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Table 4.

Pearson’s correlation coefficients for leaf traits, growth, photosynthesis, stomatal conductance, and fluorescence among all trees on August 24, 2015 (=dry).

Blue	SLA	dia gr	A	PNUE	g_s	C_i	Fv’/Fm’	ΦPSII
N	0.93^***	0.46	0.25	0.62	0.34	0.52	−0.31	0.28
SLA		0.31	−0.04	0.23	0.17	0.71	−0.40	−0.08
dia gr			−0.31	0.25	−0.39	−0.10	−0.62	0.33
A				0.60	0.92^**	0.21	0.60	0.60
PNUE					0.40	−0.10	−0.19	0.66
g_s						0.56	0.70	0.30
C_i							0.29	−0.31
Fv’/Fm’								−0.06

White
N	0.99^***	−0.58	0.06	0.14	−0.31	−0.25	0.24	−0.15
SLA		−0.54	0.12	0.13	−0.23	−0.28	0.32	−0.14
dia gr			0.52	0.47	0.70	−0.29	0.30	0.51
A				0.15	0.53	−0.92^**	0.33	0.63
PNUE					0.29	−0.04	0.08	0.15
g_s						−0.17	0.47	0.07
C_i							−0.24	−0.68
Fv’/Fm’								−0.10

Green
N	0.99^***	−0.12	−0.39	0.06	−0.26	0.06	0.32	0.16
SLA		−0.20	−0.39	0.03	−0.31	−0.03	0.29	0.21
dia gr			−0.09	−0.01	0.46	0.77^*	−0.19	−0.66
A				0.84^**	0.73^*	0.12	0.61	0.47
PNUE					0.75^*	0.29	0.80^*	0.43
g_s						0.75^*	0.44	−0.08
C_i							0.07	−0.46
Fv’/Fm’								0.32

Black
N	0.98^***	−0.40	−0.50	−0.33	−0.59	−0.17	−0.68	−0.57
SLA		−0.31	−0.73	−0.59	−0.77^*	−0.39	−0.67	−0.78^*
dia gr			−0.60	−0.42	−0.47	−0.36	0.48	−0.51
A				0.84^*	0.97^***	0.75	0.31	0.91^**
PNUE					0.85^*	0.86^*	0.56	0.58
g_s						0.87^*	0.39	0.88^**
C_i							0.28	0.44
Fv’/Fm’								0.15

Man × Black
N	0.67	−0.31	−0.48	−0.21	−0.34	−0.60	0.04	−0.02
SLA		−0.09	0.03	0.17	−0.10	−0.43	0.43	0.57
dia gr			0.65	0.41	0.61	0.67	−0.30	0.36
A				0.78^*	0.88^**	0.77^*	0.47	0.59
PNUE					0.93^**	0.78^*	0.72	0.31
g_s						0.90^**	0.50	0.30
C_i							0.21	0.12
Fv’/Fm’								0.20

Manchurian
N	0.83^**	0.74^*	0.65	0.80^*	0.34	−0.23	−0.20	0.67
SLA		0.43	0.27	0.61	0.01	−0.30	0.12	0.28
dia gr			0.71^*	0.79^*	0.64	0.10	0.13	0.48
A				0.86^**	0.61	−0.19	−0.40	0.70
PNUE					0.68	−0.04	−0.08	0.51
g_s						0.63	−0.10	0.40
C_i							0.22	−0.09
Fv’/Fm’								−0.74^*

↵*P < 0.05,
↵**P < 0.01,
↵***P < 0.001; N = leaf nitrogen (μmol mol⁻² s⁻¹); SLA = specific leaf area (m²g⁻¹); dia gr = diameter growth in 2015 (cm); A = net assimilation rate (μmol m⁻² s⁻¹); g_s = stomatal conductance (mol m⁻² s⁻¹); PNUE = photosynthetic nitrogen use efficiency (μmol mol⁻² s⁻¹); C_i = intercellular CO₂ (μmol mol⁻¹); Fv’/Fm’ = light-adapted variable fluorescence; ΦPSII = fraction of absorbed photons used by photosystem II.

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Table 5.

Pearson’s correlation coefficients for leaf traits, growth, photosynthesis, stomatal conductance, and fluorescence among all trees on September 7, 2015 (=dry).

Blue	SLA	dia gr	A	PNUE	g_s	C_i	Fv’/Fm’	ΦPSII
N	0.93***	0.46	0.31	−0.34	0.24	−0.07	0.03	−0.02
SLA		0.31	−0.01	−0.58	−0.08	−0.30	−0.27	−0.21
dia gr			0.64	0.33	0.39	−0.11	0.50	0.68
A				0.76*	0.92**	0.62	0.93***	0.74*
PNUE					0.68	0.54	0.88**	0.81*
g_s						0.85**	0.87**	0.48
C_i							0.66	0.13
Fv’/Fm’								0.71*

White
N	0.99***	−0.58	0.16	−0.77*	0.62	0.45	0.59	0.50
SLA		−0.54	0.12	−0.79*	0.63	0.48	0.64	0.45
dia gr			−0.54	0.10	−0.48	−0.15	−0.32	−0.78*
A				0.39	0.49	−0.17	0.03	0.80*
PNUE					−0.31	−0.48	−0.63	0.10
g_s						0.74*	0.56	0.40
C_i							0.42	−0.19
Fv’/Fm’								0.18

Green
N	0.99***	−0.12	−0.18	−0.78*	−0.03	0.24	−0.11	0.14
SLA		−0.20	−0.26	−0.81*	−0.06	0.33	−0.15	0.06
dia gr			0.73*	0.57	0.35	−0.53	0.22	0.64
A				0.70	0.79*	−0.22	0.80*	0.80*
PNUE					0.52	−0.20	0.50	0.38
g_s						0.40	0.76*	0.48
C_i							0.07	−0.38
Fv’/Fm’								0.65

Black
N	0.98***	−0.40	0.12	−0.50	0.20	0.29	−0.16	0.08
SLA		−0.31	0.03	−0.57	0.10	0.22	−0.16	−0.07
dia gr			0.08	0.30	0.11	0.28	0.08	0.12
A				0.75*	0.99***	−0.03	0.09	0.92**
PNUE					0.69	−0.22	0.20	0.71*
g_s						0.13	0.11	0.91**
C_i							0.14	0.03
Fv’/Fm’								−0.20

Man × Black
N	0.67	−0.31	−0.43	−0.67	−0.19	0.21	−0.02	−0.14
SLA		−0.09	−0.16	−0.39	0.44	0.68	−0.17	0.11
dia gr			−0.19	−0.14	−0.09	−0.08	−0.35	−0.58
A				0.94**	0.85**	0.13	0.26	0.74*
PNUE					0.57	−0.29	0.14	0.70
g_s						0.58	0.23	0.55
C_i							−0.14	0.18
Fv’/Fm’								−0.04

Manchurian
N	0.83**	0.74*	0.64	0.51	0.47	−0.72*	0.56	0.62
SLA		0.43	0.50	0.40	0.36	−0.67	0.71*	0.26
dia gr			0.72*	0.62	0.51	−0.71*	0.47	0.73*
A				0.98***	0.95***	−0.75*	0.70	0.88**
PNUE					0.96***	−0.70	0.67	0.87**
g_s						−0.52	0.65	0.82*
C_i							−0.57	−0.66
Fv’/Fm’								0.38

*P < 0.05,
**P < 0.01,
***P < 0.001; N = leaf nitrogen (μmol mol⁻² s⁻¹); SLA = specific leaf area (m²g⁻¹); dia gr = diameter growth in 2015 (cm); A = net assimilation rate (μmol m⁻² s⁻¹); g_s = stomatal conductance (mol m⁻² s⁻¹); PNUE = photosynthetic nitrogen use efficiency (μmol mol⁻² s⁻¹); C_i = intercellular CO₂ (μmol mol⁻¹); Fv’/Fm’ = light-adapted variable fluorescence; ΦPSII = fraction of absorbed photons used by photosystem II.