Effects of nitrogen addition level and NH4+-N to NO3--N ratio on photosynthetic characteristics and chlorophyll fluorescence parameters in Cunninghamia lanceolata seedling

doi:10.17521/cjpe.2023.0200

Abstract

Abstract:

Aims To study the effects of different nitrogen (N) addition levels and NH₄⁺-N to NO₃^--N ratios on photosynthetic characteristics and chlorophyll fluorescence parameters of Cunninghamia lanceolata will provid a scientific basis for the management of nitrogen fertilization in C. lanceolata.

Methods One-year-old seedlings of C. lanceolata were cultured in sands with three N addition levels of 0.5 (N₁), 1.0 (N₂), and 2.0 mmol·L^-1 (N₃) and seven different N form ratios (NH₄⁺-N to NO₃^--N ratios being as 10:0 (P₁), 8:2 (P₂), 6:4 (P₃), 5:5 (P₄), 4:6 (P₅), 2:8 (P₆), and 0:10 (P₇). The chlorophyll (Chl) content, photosynthetic characteristics, Chl fluorescence parameters and biomass in C. lanceolata were analyzed after 180 d treatment.

Important findings (1) The Chl a and Chl b content of C. lanceolata were the highest at P₂ and P₆ under N₁level, respectively. The Chl a, Chl b, and Chl (a+b) content were all highest at P₄ under N₂level. Both Chl a and Chl b content were the highest at P₃ among the seven ratios under N₃level. The Chl content showed an overall higher level of N₃ and N₂ than that of N₁ level. (2) The net photosynthetic rate (P_n), stomatal conductance (G_s), transpiration rate (T_r), and water use efficiency (WUE) of C. lanceolata were all higher for higher NH₄⁺-N content than for higher NO₃^--N content. The P_n was the highest at P₁ under N₃ level, but the highest at P₂ under N₂ and N₁ levels. (3) The maximum photochemical efficiency of all treatments was in the normal range of 0.80-0.85, and the potential photochemical activity of photosystem II (PSII) was significantly higher under N₃ level than that of N₁ and N₂ at P₂. The fast chlorophyll fluorescence induction curve (OJIP) of leaves deviated greatly under N₁ level at P₁. P₁, P₂, and P₄ at N₂ and P₁, P₂, and P₅ at N₃gradually decreased at the I and P phases and gradually increased at the J phase. The apparent quantum flux per unit leaf cross-sectional area, heat dissipation capacity, and the number of active reaction centers per unit leaf cross-sectional area with the increase of nitrogen content at P₂ and P₃, while the absorption flux per reaction center (RC), electron transport flux per RC, trapped energy flux per RC, and dissipated energy flux per RC at P₆ decreased significantly with the increase of nitrogen content, the differences were not significant under the other ratios. (4) Total biomass and aboveground biomass of C. lanceolata were higher for higher NH₄⁺-N content than for higher NO₃^--N content, and significantly higher under the N₂P₂ treatment than under the other treatments. At the same N form ratio, it showed that N₂ > N₃ > N₁. The root-shoot ratio of N₁ and N₃ was significantly higher than that of other treatments, and the ratio of P₆ at N₂ level was significantly higher than that of other treatments, and the ratio of P₇ at N₂ level was significantly higher than that of N₂ and N₃. The photochemical reaction efficiency of C. lanceolata seedlings could be improved and the energy utilization of the PSII reaction center could be optimized by a higher N content and a higher ammonium to nitrate ratios, which would be more beneficial for photosynthesis and biomass accumulation.

Key words: nitrogen addition, NH₄⁺-N/NO₃^--N ratio, photosynthetic characteristics, chlorophyll fluorescence, Cunninghamia lanceolata

QUAN Xiao-Qiang, WANG Yan-Ru, LI Xiao-Yu, LIANG Hai-Yan, WANG Li-Dong, YAN Xiao-Li. Effects of nitrogen addition level and NH₄⁺-N to NO₃^--N ratio on photosynthetic characteristics and chlorophyll fluorescence parameters in Cunninghamia lanceolata seedling[J]. Chin J Plant Ecol, 2024, 48(8): 1050-1064.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2023.0200

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Figures/Tables 8

References 47

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参数和公式 Parameter and formula	生物学意义 Biological meaning
F_v/F_m	最大光化学效率 Maximum photochemical efficiency
F_v/F_o	光系统II潜在光化学活性 Potential photochemical activity of photosystem II
ABS/CS_m = F_m	单位面积吸收的光能 Light energy absorbed per unit area
TR_o/CS_m = φP_o(ABS/CS_m)	单位面积捕获的光能 Light energy captured per unit area
ET_o/CS_m = φE_o(ABS/CS_m)	单位面积电子传递的量子产额 Quantum yield of electron transport per unit area
DI_o/CS_m = (ABS/CS_m) - (TR_o/CS_m)	单位面积的热耗散 Heat dissipation per unit area
RC/CS_m = φP_o × (V_J/M_o) × (ABS/CS_m)	单位面积内反应中心的数量 Number of reaction centers per unit area
ABS/RC = M_o × (1/V_j) × (1/φP_o)	单位反应中心吸收的光能 Unit of light energy absorbed by the reaction center
TR_o/RC = M_o × (1/V_j)	单位反应中心捕获的用于初级电子受体(Q_A)的能量 Unit of energy captured by the reaction center for primary electron acceptor (Q_A)
ET_o/RC = M_o × (1/V_j) × ψ_o	单位反应中心捕获的用于电子传递的能量 Unit of energy captured by a reaction center for electron transport
DI_o/RC = ABS/RC - TR_o/RC	单位反应中心耗散掉的能量 Unit of energy dissipated from the reaction center

参数和公式 Parameter and formula	生物学意义 Biological meaning
F_v/F_m	最大光化学效率 Maximum photochemical efficiency
F_v/F_o	光系统II潜在光化学活性 Potential photochemical activity of photosystem II
ABS/CS_m = F_m	单位面积吸收的光能 Light energy absorbed per unit area
TR_o/CS_m = φP_o(ABS/CS_m)	单位面积捕获的光能 Light energy captured per unit area
ET_o/CS_m = φE_o(ABS/CS_m)	单位面积电子传递的量子产额 Quantum yield of electron transport per unit area
DI_o/CS_m = (ABS/CS_m) - (TR_o/CS_m)	单位面积的热耗散 Heat dissipation per unit area
RC/CS_m = φP_o × (V_J/M_o) × (ABS/CS_m)	单位面积内反应中心的数量 Number of reaction centers per unit area
ABS/RC = M_o × (1/V_j) × (1/φP_o)	单位反应中心吸收的光能 Unit of light energy absorbed by the reaction center
TR_o/RC = M_o × (1/V_j)	单位反应中心捕获的用于初级电子受体(Q_A)的能量 Unit of energy captured by the reaction center for primary electron acceptor (Q_A)
ET_o/RC = M_o × (1/V_j) × ψ_o	单位反应中心捕获的用于电子传递的能量 Unit of energy captured by a reaction center for electron transport
DI_o/RC = ABS/RC - TR_o/RC	单位反应中心耗散掉的能量 Unit of energy dissipated from the reaction center

参数 Parameter	氮添加水平 Nitrogen addition level (N)		铵硝态氮配比 NH₄⁺-N/NO₃^--N ratio (R)		氮添加水平×铵硝态氮配比 N × R
参数 Parameter	F	p	F	p	F	p
叶绿素a含量 Chlorophyll a content	4.47	<0.05	3.34	<0.01	6.99	<0.01
叶绿素b含量 Chlorophyll b content	3.83	<0.05	1.34	0.25	5.17	<0.01
叶绿素(a+b)含量 Chlorophyll (a+b) content	2.85	0.07	1.57	0.17	2.95	<0.01
净光合速率 Net photosynthetic rate	2.65	0.07	54.21	<0.01	8.42	<0.01
蒸腾速率 Transpiration rate	28.29	<0.01	50.74	<0.01	6.67	<0.01
气孔导度 Stomatal conductance	62.30	<0.01	33.68	<0.01	15.57	<0.01
水分利用效率 Water use efficiency	501.75	<0.01	31.75	<0.01	31.36	<0.01
最大光化学效率 Maximum photochemical efficiency	5.87	<0.01	3.35	<0.01	0.67	0.78
光系统II潜在化学活性 Potential chemical activity of photosystem II	6.90	<0.01	3.75	<0.01	0.81	0.64
地上生物量 Above-ground biomass	21.45	<0.01	18.33	<0.01	2.07	0.04
地下生物量 Below-ground biomass	17.22	<0.01	20.37	<0.01	1.33	0.24
总生物量 Total biomass	22.18	<0.01	9.47	<0.01	1.85	0.07
根冠比 Root-shoot ratio	19.65	<0.01	31.18	<0.01	1.64	0.12

参数 Parameter	氮添加水平 Nitrogen addition level (N)		铵硝态氮配比 NH₄⁺-N/NO₃^--N ratio (R)		氮添加水平×铵硝态氮配比 N × R
参数 Parameter	F	p	F	p	F	p
叶绿素a含量 Chlorophyll a content	4.47	<0.05	3.34	<0.01	6.99	<0.01
叶绿素b含量 Chlorophyll b content	3.83	<0.05	1.34	0.25	5.17	<0.01
叶绿素(a+b)含量 Chlorophyll (a+b) content	2.85	0.07	1.57	0.17	2.95	<0.01
净光合速率 Net photosynthetic rate	2.65	0.07	54.21	<0.01	8.42	<0.01
蒸腾速率 Transpiration rate	28.29	<0.01	50.74	<0.01	6.67	<0.01
气孔导度 Stomatal conductance	62.30	<0.01	33.68	<0.01	15.57	<0.01
水分利用效率 Water use efficiency	501.75	<0.01	31.75	<0.01	31.36	<0.01
最大光化学效率 Maximum photochemical efficiency	5.87	<0.01	3.35	<0.01	0.67	0.78
光系统II潜在化学活性 Potential chemical activity of photosystem II	6.90	<0.01	3.75	<0.01	0.81	0.64
地上生物量 Above-ground biomass	21.45	<0.01	18.33	<0.01	2.07	0.04
地下生物量 Below-ground biomass	17.22	<0.01	20.37	<0.01	1.33	0.24
总生物量 Total biomass	22.18	<0.01	9.47	<0.01	1.85	0.07
根冠比 Root-shoot ratio	19.65	<0.01	31.18	<0.01	1.64	0.12

指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	氮添加水平 Nitrogen addition level
指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	N₁	N₂	N₃
叶绿素a含量 Chlorophyll a content (mg·g^-1)	P₁	0.374 ± 0.026^ABCa	0.367 ± 0.011^Ba	0.414 ± 0.016^ABCa
	P₂	0.418 ± 0.019^Aa	0.333 ± 0.015^Bb	0.436 ± 0.020^ABa
	P₃	0.372 ± 0.013^ABCb	0.342 ± 0.033^Bb	0.457 ± 0.010^Aa
	P₄	0.315 ± 0.015^Cc	0.488 ± 0.002^Aa	0.410 ± 0.020^ABCb
	P₅	0.371 ± 0.016^ABCa	0.410 ± 0.038^ABa	0.332 ± 0.023^Ca
	P₆	0.402 ± 0.017^ABa	0.368 ± 0.004^Ba	0.379 ± 0.019^ABCa
	P₇	0.327 ± 0.016^BCa	0.343 ± 0.009^Ba	0.359 ± 0.020^BCa
叶绿素b含量 Chlorophyll b content (mg·g^-1)	P₁	0.215 ± 0.009^ABb	0.245 ± 0.006^ABa	0.254 ± 0.003^ABa
	P₂	0.253 ± 0.012^Aa	0.207 ± 0.008^Bb	0.246 ± 0.013^ABab
	P₃	0.235 ± 0.006^ABab	0.223 ± 0.013^Bb	0.265 ± 0.003^Aa
	P₄	0.216 ± 0.012^ABb	0.289 ± 0.008^Aa	0.237 ± 0.011^ABb
	P₅	0.226 ± 0.008^ABab	0.248 ± 0.014^ABa	0.206 ± 0.003^Bb
	P₆	0.254 ± 0.007^Aa	0.227 ± 0.007^Ba	0.260 ± 0.018^ABa
	P₇	0.208 ± 0.007^Ba	0.247 ± 0.017^ABa	0.250 ± 0.021^ABa
叶绿素(a+b)含量 Chlorophyll (a+b) content (mg·g^-1)	P₁	0.624 ± 0.058^Aa	0.612 ± 0.017^Ba	0.667 ± 0.018^Aa
	P₂	0.619 ± 0.060^Aa	0.507 ± 0.035^Ba	0.617 ± 0.072^Aa
	P₃	0.628 ± 0.030^Aab	0.565 ± 0.046^Bb	0.723 ± 0.012^Aa
	P₄	0.531 ± 0.024^Ab	0.803 ± 0.033^Aa	0.648 ± 0.030^Ab
	P₅	0.597 ± 0.024^Aa	0.657 ± 0.052^ABa	0.552 ± 0.033^Aa
	P₆	0.656 ± 0.022^Aa	0.646 ± 0.030^ABa	0.684 ± 0.066^Aa
	P₇	0.514 ± 0.032^Aa	0.629 ± 0.056^ABa	0.660 ± 0.076^Aa

Effects of nitrogen addition level and NH₄⁺-N to NO₃^--N ratio on photosynthetic characteristics and chlorophyll fluorescence parameters in Cunninghamia lanceolata seedling

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指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	氮添加水平 Nitrogen addition level
指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	N₁	N₂	N₃
地上生物量 Above-ground biomass (g·tree^-1)	P₁	4.00 ± 0.25^Aa	6.01 ± 0.88^ABCa	5.76 ± 0.34^Aa
	P₂	3.82 ± 0.19^Ab	7.17 ± 0.36^Aa	5.28 ± 0.69^ABab
	P₃	3.99 ± 0.08^Ab	6.52 ± 0.22^ABa	5.12 ± 0.13^ABab
	P₄	3.42 ± 0.61^Aa	4.61 ± 0.51^BCDa	4.11 ± 0.22^ABa
	P₅	2.96 ± 0.45^Ab	3.74 ± 0.34^CDa	3.57 ± 0.17^Bab
	P₆	2.61 ± 0.19^Aa	3.26 ± 0.32^Da	3.35 ± 0.41^Ba
	P₇	2.95 ± 0.15^Aa	2.91 ± 0.60^Da	3.27 ± 0.74^Ba
地下生物量 Below-ground biomass (g·tree^-1)	P₁	1.30 ± 0.10^Ca	1.18 ± 0.17^CDa	1.07 ± 0.10^BCa
	P₂	1.16 ± 0.18^Ca	1.07 ± 0.17^Da	1.01 ± 0.07^BCa
	P₃	1.35 ± 0.10^BCa	1.33 ± 0.14^BCDa	0.96 ± 0.03^Ca
	P₄	1.58 ± 0.10^ABCa	1.46 ± 0.17^BCDa	1.33 ± 0.05^ABCa
	P₅	1.90 ± 0.10^ABa	1.86 ± 0.16^ABCa	1.20 ± 0.12^ABCb
	P₆	2.11 ± 0.01^Aa	2.20 ± 0.10^Aa	1.63 ± 0.05^ABa
	P₇	1.67 ± 0.14^ABCb	2.03 ± 0.13^ABa	1.47 ± 0.23^ABb
总生物量 Total biomass (g·tree^-1)	P₁	5.29 ± 0.30^Aa	7.19 ± 0.80^ABCa	6.83 ± 0.34^Aa
	P₂	4.98 ± 0.22^Ab	8.24 ± 0.23^Aa	6.29 ± 0.64^Aab
	P₃	5.34 ± 0.18^Ab	7.85 ± 0.10^ABa	6.09 ± 0.15^Aab
	P₄	5.00 ± 0.53^Aa	6.07 ± 0.68^ABCa	5.43 ± 0.26^Aa
	P₅	4.85 ± 0.50^Aa	5.60 ± 0.26^BCa	4.78 ± 0.23^Aa
	P₆	4.72 ± 0.18^Aa	5.46 ± 0.43^BCa	4.99 ± 0.36^Aa
	P₇	4.62 ± 0.29^Aa	4.95 ± 0.69^Ca	4.74 ± 0.53^Aa
根冠比 Root-shoot ratio	P₁	0.33 ± 0.04^Ba	0.21 ± 0.07^Ca	0.19 ± 0.03^Ba
	P₂	0.31 ± 0.01^Ba	0.16 ± 0.03^Cb	0.19 ± 0.02^Bb
	P₃	0.35 ± 0.07^Ba	0.20 ± 0.01^Ca	0.19 ± 0.02^Ba
	P₄	0.46 ± 0.02^Ba	0.32 ± 0.03^BCa	0.34 ± 0.07^ABa
	P₅	0.64 ± 0.02^ABa	0.50 ± 0.06^ABab	0.34 ± 0.02^ABb
	P₆	0.85 ± 0.15^Aa	0.68 ± 0.04^Aa	0.50 ± 0.07^Aa
	P₇	0.59 ± 0.09^ABa	0.70 ± 0.03^Aa	0.45 ± 0.04^Aa

指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	氮添加水平 Nitrogen addition level
指标 Index	铵硝态氮配比 NH₄⁺-N to NO₃^--N ratio	N₁	N₂	N₃
地上生物量 Above-ground biomass (g·tree^-1)	P₁	4.00 ± 0.25^Aa	6.01 ± 0.88^ABCa	5.76 ± 0.34^Aa
	P₂	3.82 ± 0.19^Ab	7.17 ± 0.36^Aa	5.28 ± 0.69^ABab
	P₃	3.99 ± 0.08^Ab	6.52 ± 0.22^ABa	5.12 ± 0.13^ABab
	P₄	3.42 ± 0.61^Aa	4.61 ± 0.51^BCDa	4.11 ± 0.22^ABa
	P₅	2.96 ± 0.45^Ab	3.74 ± 0.34^CDa	3.57 ± 0.17^Bab
	P₆	2.61 ± 0.19^Aa	3.26 ± 0.32^Da	3.35 ± 0.41^Ba
	P₇	2.95 ± 0.15^Aa	2.91 ± 0.60^Da	3.27 ± 0.74^Ba
地下生物量 Below-ground biomass (g·tree^-1)	P₁	1.30 ± 0.10^Ca	1.18 ± 0.17^CDa	1.07 ± 0.10^BCa
	P₂	1.16 ± 0.18^Ca	1.07 ± 0.17^Da	1.01 ± 0.07^BCa
	P₃	1.35 ± 0.10^BCa	1.33 ± 0.14^BCDa	0.96 ± 0.03^Ca
	P₄	1.58 ± 0.10^ABCa	1.46 ± 0.17^BCDa	1.33 ± 0.05^ABCa
	P₅	1.90 ± 0.10^ABa	1.86 ± 0.16^ABCa	1.20 ± 0.12^ABCb
	P₆	2.11 ± 0.01^Aa	2.20 ± 0.10^Aa	1.63 ± 0.05^ABa
	P₇	1.67 ± 0.14^ABCb	2.03 ± 0.13^ABa	1.47 ± 0.23^ABb
总生物量 Total biomass (g·tree^-1)	P₁	5.29 ± 0.30^Aa	7.19 ± 0.80^ABCa	6.83 ± 0.34^Aa
	P₂	4.98 ± 0.22^Ab	8.24 ± 0.23^Aa	6.29 ± 0.64^Aab
	P₃	5.34 ± 0.18^Ab	7.85 ± 0.10^ABa	6.09 ± 0.15^Aab
	P₄	5.00 ± 0.53^Aa	6.07 ± 0.68^ABCa	5.43 ± 0.26^Aa
	P₅	4.85 ± 0.50^Aa	5.60 ± 0.26^BCa	4.78 ± 0.23^Aa
	P₆	4.72 ± 0.18^Aa	5.46 ± 0.43^BCa	4.99 ± 0.36^Aa
	P₇	4.62 ± 0.29^Aa	4.95 ± 0.69^Ca	4.74 ± 0.53^Aa
根冠比 Root-shoot ratio	P₁	0.33 ± 0.04^Ba	0.21 ± 0.07^Ca	0.19 ± 0.03^Ba
	P₂	0.31 ± 0.01^Ba	0.16 ± 0.03^Cb	0.19 ± 0.02^Bb
	P₃	0.35 ± 0.07^Ba	0.20 ± 0.01^Ca	0.19 ± 0.02^Ba
	P₄	0.46 ± 0.02^Ba	0.32 ± 0.03^BCa	0.34 ± 0.07^ABa
	P₅	0.64 ± 0.02^ABa	0.50 ± 0.06^ABab	0.34 ± 0.02^ABb
	P₆	0.85 ± 0.15^Aa	0.68 ± 0.04^Aa	0.50 ± 0.07^Aa
	P₇	0.59 ± 0.09^ABa	0.70 ± 0.03^Aa	0.45 ± 0.04^Aa