塔克拉玛干沙漠南缘豆科与非豆科植物的氮分配

doi:10.3773/j.issn.1005-264x.2010.09.003

摘要/Abstract

摘要：

在豆科与非豆科植物光合特性的研究中发现, 非豆科植物具有更高的光合速率, 与其低的叶氮含量相矛盾。在沙漠中氮素是限制植物生长的关键因子之一, 考虑到豆科植物的生物固氮作用和叶氮大部分分配于光合系统, 我们假设: (1)非豆科植物具有更低的叶氮含量; (2)分配更少的叶氮于光合系统; (3)具有更高的最大净光合速率(P_max)和光合氮素利用效率(PNUE)。为了验证这些假设, 以塔克拉玛干沙漠南缘的豆科植物骆驼刺(Alhagi sparsifolia)和非豆科植物柽柳(Tamarix ramosissima)、花花柴(Karelinia caspica)为研究对象, 比较了它们的叶氮含量、氮分配、P_max和PNUE等。结果表明: (1)非豆科植物比豆科植物确实有更低的叶氮含量, 且差异达到显著水平; (2)非豆科植物分配更少的叶氮于光合系统, 但在光合系统内部具有更高效的氮分配机制; (3)非豆科植物具有更高的P_max和PNUE。在光合系统内部, 非豆科植物分配更多的叶氮于羧化系统, 而豆科植物分配更多的叶氮于捕光系统。对于非豆科植物而言, 其更高的P_max、PNUE、水分利用效率和表观量子产量, 取决于将更多的叶氮投入到羧化和电子传递系统中。这些生理优势决定了塔克拉玛干沙漠南缘非豆科植物高效的资源捕捉和利用能力。

关键词: 豆科植物, 氮分配, 非豆科植物, 光合氮素利用效率, 塔克拉玛干沙漠

Abstract:

Aims We previously found that non-leguminous species had a higher photosynthetic rate despite lower leaf N content. Nitrogen is a critical factor limiting plant growth in the desert. Our objectives were to determine if, as hypothesized, non-leguminous species have lower leaf N content, allocate a lower fraction of leaf N to photosynthesis and have higher maximum net photosynthetic rate (P_max) and photosynthetic N-use efficiency (PNUE).

Methods We compared the leguminous species Alhagi sparsifolia and non-leguminous species Karelinia caspica and Tamarix ramosissima in their typical habitat at the southern fringe of China’s Taklamakan Desert.

Important findings As hypothesized, the non-leguminous species had significantly lower leaf N content and allocated a lower fraction of leaf N to photosynthesis. They also were more efficient in photosynthetic N partitioning. The non-leguminous species partitioned a higher fraction of the photosynthetic N to carboxylation and showed higher use efficiency of the photosynthetic N, while the leguminous species partitioned a higher fraction of the photosynthetic N to light-harvesting components. For the non-leguminous species, the higher fraction of leaf N allocated to carboxylation and bioenergetics led to higher P_max and therefore to higher PNUE, water-use efficiency and apparent quantum yield. These physiological advantages of the non-leguminous species and their higher leaf area ratio may contribute to their higher resource capture ability.

Key words: legume species, nitrogen allocation, non-legume species, photosynthetic nitrogen-use efficiency, Taklamakan Desert

朱军涛, 李向义, 张希明, 林丽莎, 杨尚功. 塔克拉玛干沙漠南缘豆科与非豆科植物的氮分配. 植物生态学报, 2010, 34(9): 1025-1032. DOI: 10.3773/j.issn.1005-264x.2010.09.003

ZHU Jun-Tao, LI Xiang-Yi, ZHANG Xi-Ming, LIN Li-Sha, YANG Shang-Gong. Nitrogen allocation and partitioning within a leguminous and two non-leguminous plant species growing at the southern fringe of China’s Taklamakan Desert. Chinese Journal of Plant Ecology, 2010, 34(9): 1025-1032. DOI: 10.3773/j.issn.1005-264x.2010.09.003

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图/表 4

参考文献 37

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变量 Variables	花花柴 Karelinia caspica	柽柳 Tamarix ramosissima	骆驼刺 Alhagi sparsifolia	F值 F value
表观量子产额 Apparent quantum requirement (AQY) (mol·mol^-1)	0.038 ± 0.003^a	0.034 ± 0.004^a	0.029 ± 0.002^b	3.625^*
最大净光合速率 Maximum net photosynthetic rate (P_max) (μmol·m^-2·s^-1)	11.7 ± 2.3^a	9.5 ± 3.0^a	6.9 ± 1.3^b	12.724^**
最大电子传递速率 Maximum electron transport rate (J_max) (μmol·m^-2·s^-1)	105.2 ± 10.1^a	94.8 ± 7.5^b	87.8 ± 6.3^b	5.615^*
最大羧化速率 Maximum carboxylation efficiency (V_cmax) (μmol·m^-2·s^-1)	62.5 ± 3.7^a	38.7 ± 4.6^b	30.8 ± 2.5^c	10.082^**
叶氮在羧化系统的比例 The fraction of the total leaf nitrogen allocated to carboxylation (P_C) (g·g^-1)	0.23 ± 0.04^a	0.11 ± 0.03^b	0.07 ± 0.02^c	5.579^***
叶氮在生物力能学组分的比例 The fraction of the total leaf nitrogen allocated to bioenergetics (P_B) (g·g^-1)	0.04 ± 0.003	0.03 ± 0.003	0.03 ± 0.002	3.832
P_C、P_B之和 The sum of P_C and P_B (P_{C + B}) (g·g^-1)	0.27 ± 0.05^a	0.14 ± 0.03^b	0.10 ± 0.02^c	6.215^***
叶氮在捕光系统的比例 The fraction of the total leaf nitrogen allocated to light-harvesting components (P_L) (g·g^-1)	0.16 ± 0.02^b	0.20 ± 0.02^b	0.29 ± 0.04^a	8.232^**
叶氮在光合系统的比例 The fraction of the total leaf nitrogen allocated to all components of the photosynthetic apparatus (P_T) (g·g^-1)	0.43 ± 0.03^a	0.39 ± 0.05^b	0.38 ± 0.05^b	10.253^**
叶氮在羧化系统的数量 Nitrogen content in carboxylation (N_C) (g·m^-2)	0.48 ± 0.04^a	0.31 ± 0.02^b	0.23 ± 0.03^c	12.325^***
叶氮在生物力能学组分的数量 Nitrogen content in bioenergetics (N_B) (g·m^-2)	0.08 ± 0.005	0.08 ± 0.003	0.07 ± 0.005	2.323
N_C、N_B之和 The sum of N_Cand N_B (N_{C + B}) (g·m^-2)	0.55 ± 0.03^a	0.39 ± 0.02^b	0.31 ± 0.03^c	14.536^***
叶氮在捕光系统的数量 Nitrogen content in light-harvesting components (N_L) (g·m^-2)	0.43 ± 0.02^b	0.56 ± 0.03^b	0.97 ± 0.02^a	8.752^***
叶氮在光合系统的数量 Nitrogen content in all components of the photosynthetic apparatus (N_P) (g·m^-2)	0.89 ± 0.06^b	0.96 ± 0.05^b	1.27 ± 0.07^a	5.672^**
N_C、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to carboxylation (N_C/N_P)	0.54 ± 0.04^a	0.32 ± 0.05^b	0.18 ± 0.07^c	10.517^***
N_B、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to bioenergetics (N_B/N_P)	0.09 ± 0.01^a	0.08 ± 0.01^a	0.06 ± 0.01^b	4.265^*
N_L、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to light-harvesting components (N_L/N_P)	0.37 ± 0.02^c	0.58 ± 0.04^b	0.76 ± 0.05^a	13.360^***
单位面积叶氮含量 Leaf nitrogen content per area (N_A) (g·m^-2)	2.07 ± 0.10^c	2.81 ± 0.09^b	3.33 ± 0.13^a	9.226^**
光合氮素利用效率 Photosynthetic nitrogen-use efficiency (PNUE) (μmol·g^-1·s^-1)	9.50 ± 0.93^a	8.89 ± 0.75^a	5.82 ± 0.28^b	6.054^**
P_max、N_P 之比 Photosynthetic-use efficiency of the photosynthetic nitrogen (P_max/N_P) (μmol·g^-1·s^-1)	13.15 ± 1.55^a	9.86 ± 1.24^b	5.43 ± 1.15^c	4.835^***
气孔导度 Stomatal conductance (G_s) (mol·m^-2·s^-1)	0.26 ± 0.02	0.25 ± 0.02	0.24 ± 0.03	3.324
胞间 CO₂ 浓度 Intercellular CO₂concentration (C_i) (μmol·mol^-1)	220.5 ± 7.2	230.6 ± 5.5	237.8 ± 7.0	4.458
叶绿素含量 Leaf chlorophyll content (Chl) (μmol·m^-2)	0.24 ± 0.04^b	0.22 ± 0.03^b	0.46 ± 0.05^a	5.725^**
叶生物量分数 Leaf mass fraction (LMF) (g·g^-1)	0.65 ± 0.03^a	0.58 ± 0.04^a	0.35 ± 0.03^b	16.535^***
叶面积比 Leaf area ratio (LAR) (cm²·g^-1)	97.5 ± 5.8^a	88.7 ± 7.5^a	58.5 ± 5.5^b	10.336^**
比叶面积 Specfic leaf area (SLA) (cm²·g^-1)	153.4 ± 18.3^a	145.2 ± 22.7^b	141.5 ± 17.8^b	2.415^**

变量 Variables	花花柴 Karelinia caspica	柽柳 Tamarix ramosissima	骆驼刺 Alhagi sparsifolia	F值 F value
表观量子产额 Apparent quantum requirement (AQY) (mol·mol^-1)	0.038 ± 0.003^a	0.034 ± 0.004^a	0.029 ± 0.002^b	3.625^*
最大净光合速率 Maximum net photosynthetic rate (P_max) (μmol·m^-2·s^-1)	11.7 ± 2.3^a	9.5 ± 3.0^a	6.9 ± 1.3^b	12.724^**
最大电子传递速率 Maximum electron transport rate (J_max) (μmol·m^-2·s^-1)	105.2 ± 10.1^a	94.8 ± 7.5^b	87.8 ± 6.3^b	5.615^*
最大羧化速率 Maximum carboxylation efficiency (V_cmax) (μmol·m^-2·s^-1)	62.5 ± 3.7^a	38.7 ± 4.6^b	30.8 ± 2.5^c	10.082^**
叶氮在羧化系统的比例 The fraction of the total leaf nitrogen allocated to carboxylation (P_C) (g·g^-1)	0.23 ± 0.04^a	0.11 ± 0.03^b	0.07 ± 0.02^c	5.579^***
叶氮在生物力能学组分的比例 The fraction of the total leaf nitrogen allocated to bioenergetics (P_B) (g·g^-1)	0.04 ± 0.003	0.03 ± 0.003	0.03 ± 0.002	3.832
P_C、P_B之和 The sum of P_C and P_B (P_{C + B}) (g·g^-1)	0.27 ± 0.05^a	0.14 ± 0.03^b	0.10 ± 0.02^c	6.215^***
叶氮在捕光系统的比例 The fraction of the total leaf nitrogen allocated to light-harvesting components (P_L) (g·g^-1)	0.16 ± 0.02^b	0.20 ± 0.02^b	0.29 ± 0.04^a	8.232^**
叶氮在光合系统的比例 The fraction of the total leaf nitrogen allocated to all components of the photosynthetic apparatus (P_T) (g·g^-1)	0.43 ± 0.03^a	0.39 ± 0.05^b	0.38 ± 0.05^b	10.253^**
叶氮在羧化系统的数量 Nitrogen content in carboxylation (N_C) (g·m^-2)	0.48 ± 0.04^a	0.31 ± 0.02^b	0.23 ± 0.03^c	12.325^***
叶氮在生物力能学组分的数量 Nitrogen content in bioenergetics (N_B) (g·m^-2)	0.08 ± 0.005	0.08 ± 0.003	0.07 ± 0.005	2.323
N_C、N_B之和 The sum of N_Cand N_B (N_{C + B}) (g·m^-2)	0.55 ± 0.03^a	0.39 ± 0.02^b	0.31 ± 0.03^c	14.536^***
叶氮在捕光系统的数量 Nitrogen content in light-harvesting components (N_L) (g·m^-2)	0.43 ± 0.02^b	0.56 ± 0.03^b	0.97 ± 0.02^a	8.752^***
叶氮在光合系统的数量 Nitrogen content in all components of the photosynthetic apparatus (N_P) (g·m^-2)	0.89 ± 0.06^b	0.96 ± 0.05^b	1.27 ± 0.07^a	5.672^**
N_C、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to carboxylation (N_C/N_P)	0.54 ± 0.04^a	0.32 ± 0.05^b	0.18 ± 0.07^c	10.517^***
N_B、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to bioenergetics (N_B/N_P)	0.09 ± 0.01^a	0.08 ± 0.01^a	0.06 ± 0.01^b	4.265^*
N_L、N_P 之比 The fraction of the photosynthetic nitrogen partitioned to light-harvesting components (N_L/N_P)	0.37 ± 0.02^c	0.58 ± 0.04^b	0.76 ± 0.05^a	13.360^***
单位面积叶氮含量 Leaf nitrogen content per area (N_A) (g·m^-2)	2.07 ± 0.10^c	2.81 ± 0.09^b	3.33 ± 0.13^a	9.226^**
光合氮素利用效率 Photosynthetic nitrogen-use efficiency (PNUE) (μmol·g^-1·s^-1)	9.50 ± 0.93^a	8.89 ± 0.75^a	5.82 ± 0.28^b	6.054^**
P_max、N_P 之比 Photosynthetic-use efficiency of the photosynthetic nitrogen (P_max/N_P) (μmol·g^-1·s^-1)	13.15 ± 1.55^a	9.86 ± 1.24^b	5.43 ± 1.15^c	4.835^***
气孔导度 Stomatal conductance (G_s) (mol·m^-2·s^-1)	0.26 ± 0.02	0.25 ± 0.02	0.24 ± 0.03	3.324
胞间 CO₂ 浓度 Intercellular CO₂concentration (C_i) (μmol·mol^-1)	220.5 ± 7.2	230.6 ± 5.5	237.8 ± 7.0	4.458
叶绿素含量 Leaf chlorophyll content (Chl) (μmol·m^-2)	0.24 ± 0.04^b	0.22 ± 0.03^b	0.46 ± 0.05^a	5.725^**
叶生物量分数 Leaf mass fraction (LMF) (g·g^-1)	0.65 ± 0.03^a	0.58 ± 0.04^a	0.35 ± 0.03^b	16.535^***
叶面积比 Leaf area ratio (LAR) (cm²·g^-1)	97.5 ± 5.8^a	88.7 ± 7.5^a	58.5 ± 5.5^b	10.336^**
比叶面积 Specfic leaf area (SLA) (cm²·g^-1)	153.4 ± 18.3^a	145.2 ± 22.7^b	141.5 ± 17.8^b	2.415^**