杉木幼苗生物量分配格局对氮添加的响应

doi:10.17521/cjpe.2021.0135

植物生态学报 ›› 2021, Vol. 45 ›› Issue (11): 1231-1240.DOI: 10.17521/cjpe.2021.0135 cstr: 32100.14.cjpe.2021.0135

杉木幼苗生物量分配格局对氮添加的响应

王娇¹^,³, 关欣¹^,², 张伟东¹^,², 黄苛¹^,², 朱睦楠¹^,², 杨庆朋¹^,²^,^*()

¹中国科学院会同森林生态实验站, 中国科学院森林生态与管理重点实验室, 中国科学院沈阳应用生态研究所, 沈阳 110016
²亚热带森林生态系统结构与服务功能湖南省重点实验室, 湖南会同 418307
³中国科学院大学资源与环境学院, 北京 101408

收稿日期:2021-04-09 接受日期:2021-07-08 出版日期:2021-11-20 发布日期:2021-09-18
作者简介:* (yqp226@iae.ac.cn)
基金资助:
国家自然科学基金(41630755);国家自然科学基金(41977092);国家自然科学基金(41877092)

Responses of biomass allocation patterns to nitrogen addition of Cunninghamia lanceolata seedlings

WANG Jiao¹^,³, GUAN Xin¹^,², ZHANG Wei-Dong¹^,², HUANG Ke¹^,², ZHU Mu-Nan¹^,², YANG Qing-Peng¹^,²^,^*()

¹Huitong Experimental Station of Forest Ecology, Key Laboratory of Forest Ecology and Management, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
²Hunan Key Laboratory for Structure and Ecosystem Service of Subtropical Forest, Huitong, Hunan 418307, China
³College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 101408, China

Received:2021-04-09 Accepted:2021-07-08 Online:2021-11-20 Published:2021-09-18
Supported by:
National Natural Science Foundation of China(41630755);National Natural Science Foundation of China(41977092);National Natural Science Foundation of China(41877092)

摘要/Abstract

摘要：

大气氮(N)沉降的急剧增加可能会对植物碳(C)固定和分配产生深远影响。然而, N添加如何影响碳水化合物在植物不同器官之间的分配动态并不十分清楚。该研究利用杉木(Cunninghamia lanceolata)幼苗盆栽试验, 设置N添加处理, 测定分析幼苗非结构性碳水化合物(NSC)与结构性碳水化合物(SC)含量和库的变化, 以探讨N添加后杉木幼苗不同器官中NSC与SC的分配模式及调控机制。结果发现: (1) N添加虽然显著增加叶片净光合速率(143.96%), 但却降低了叶片中的NSC含量和库; N添加导致一年生茎的淀粉含量显著下降, 而可溶性糖含量的变化不显著, 当年生茎的NSC组分含量和库没有显著变化; 幼苗根系的NSC及其组分含量和库也有降低的趋势。(2) N添加后地下与地上生物量的比值降低22.09%, 其中SC库比值降低31.07%, 而NSC库比值无显著变化。(3) N添加使地上部分的磷(P)库显著增加, 使地下与地上P库的比值降低了57.02%, 而N库的比值无显著变化。(4) N添加后土壤pH由(4.94 ± 0.09)显著降低到(4.02 ± 0.04), 铵态N和硝态N含量分别增加7.17倍和11.55倍, 土壤有效P含量也增加了42.86%, 而土壤中脲酶(62.75%)和酸性磷酸酶(56.52%)的活性显著降低。研究表明, 低养分条件下杉木幼苗主要通过构建根系结构增加养分吸收, 而非通过向根系分配更多的NSC, 而N添加驱动的养分缓解使更多的碳水化合物分配到地上器官, 导致地上部分SC积累。

关键词: 氮添加, 生物量分配, 碳水化合物, 非结构性碳, 磷, 最优分配理论

Abstract:

Aims Sharply increasing atmospheric nitrogen (N) deposition may have profound effects on carbon (C) fixation and allocation of plants. However, a comprehensive understanding of how nitrogen addition influences allocation dynamics of carbohydrates among different organs in plants is still lacking.
Methods In this study, a pot experiment of seedlings of Cunninghamia lanceolate was established to investigate the allocation pattern and regulation mechanism of non-structural carbohydrates (NSC) and structural carbohydrates (SC) in different organs of C. lanceolate seedlings after N addition. The concentrations and pool sizes of NSC and SC were measured.
Important findings The results showed that: (1) N addition significantly increased the net photosynthetic rate by 143.96%, but decreased the concentration and pool size of NSC in needles. There was no significant change in the components of NSC concentration and pool in current-year stems. Furthermore, N addition resulted in a significant decrease in the starch concentration of one-year-old stems, but a non-significant change in soluble sugar concentration. In addition, the components of NSC concentration and pool in seedlings roots also tended to decrease. (2) After N addition, the ratio of below- to above-ground biomass decreased by 22.09%, and the ratio of below- to above-ground SC pool also decreased by 31.07%, and the ratio of below- to above-ground NSC pool showed no significant difference between control and N addition treatment. (3) N addition significantly increased the phosphorus (P) pool size in the above-ground part and decreased the ratio of below- to above-ground P pool, while the ratio of below- to above-ground N pool showed no obvious difference between control and N addition treatment. (4) Under N addition, soil pH decreased significantly from 4.94 to 4.02, ammonium and nitrate nitrogen concentration increased by 7.17 times and 11.55 times, respectively, and soil available P concentration increased by 42.86%, while the activities of urease (62.75%) and acid phosphatase (56.52%) in the soil decreased significantly compared to control. Our results indicates that C. lanceolate seedlings increased nutrient uptake amount mainly by the construction of root structure, but not by the allocation of more NSC to the root under low nutrient conditions, while nutrient alleviation driven by nitrogen addition resulted in more carbohydrates allocation to above-ground organs, resulting in the accumulation of SC in above-ground parts.

Key words: nitrogen addition, biomass allocation, carbohydrates, non-structural carbohydrates, phosphorus, optimal partitioning theory

王娇, 关欣, 张伟东, 黄苛, 朱睦楠, 杨庆朋. 杉木幼苗生物量分配格局对氮添加的响应. 植物生态学报, 2021, 45(11): 1231-1240. DOI: 10.17521/cjpe.2021.0135

WANG Jiao, GUAN Xin, ZHANG Wei-Dong, HUANG Ke, ZHU Mu-Nan, YANG Qing-Peng. Responses of biomass allocation patterns to nitrogen addition of Cunninghamia lanceolata seedlings. Chinese Journal of Plant Ecology, 2021, 45(11): 1231-1240. DOI: 10.17521/cjpe.2021.0135

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0135

https://www.plant-ecology.com/CN/Y2021/V45/I11/1231

图/表 9

图1 氮添加对杉木幼苗净光合速率(A)、生物量(B)及不同器官(O)生物量(C)的影响(平均值±标准误, n = 5)。CK, 对照, 不添加氮肥; N, 添加氮肥处理。CN, 当年生叶; CR, 粗根; CS, 当年生茎; FR, 细根; ON, 一年生叶; OS, 一年生茎。图中不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 1 Effect of nitrogen addition on net photosynthetic rate (A), total plant biomass (B) and biomass of different organs (O) (C) of Cunninghamia lanceolate seedlings (mean ± SE, n = 5). CK, control, no nitrogen fertilizer addition; N, nitrogen addition. CN, current-year needles; CR, coarse roots; CS, current-year stems; FR, fine roots; ON, one-year-old needles; OS, one-year- old stems. Different lowercase letters indicate significant difference between treatments (p < 0.05).

图2 氮添加对杉木幼苗不同器官(O)非结构性碳水化合物(NSC)含量(A)、可溶性糖含量(B)、淀粉(C)含量、NSC库(D)、可溶性糖库(E)和淀粉库(F)的影响(平均值±标准误, n = 5)。CN, 当年生叶; CR, 粗根; CS, 当年生茎; FR, 细根; ON, 一年生叶; OS, 一年生茎。CK, 对照, 不添加氮肥; N, 添加氮肥处理。图中不同小写字母表示同一器官内不同处理间差异显著(p < 0.05)。

Fig. 2 Effect of nitrogen addition on the concentration of non-structural carbohydrates (NSC)(A), soluble sugar (B) starch (C), and the pool sizes of non-structural carbohydrates (NSC)(D), soluble sugar (E) and starch (F) in different organs (O) of Cunninghamia lanceolate seedlings (mean ± SE, n = 5). CN, current-year needles; CR, coarse roots; CS, current-year stems; FR, fine roots; ON, one-year-old needles; OS, one-year-old stems. CK, control, no nitrogen fertilizer addition; N, nitrogen addition. The different lowercase letters of the same organs indicate significant difference between treatments (p < 0.05).

表1 氮(N)添加对杉木幼苗各器官非结构性碳水化合物(NSC)、可溶性糖、淀粉、N和磷(P)含量的影响

Table 1 Effect of nitrogen (N) addition on the concentrations of non-structural carbohydrates (NSC), soluble sugar, starch, N and phosphorus (P) in different organs of Cunninghamia lanceolate seedlings

器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

图3 氮添加对杉木幼苗不同器官(O)氮(N)含量(A)、磷(P)含量(B)、N库(C)和P库(D)的影响(平均值±标准误, n = 5)。CN, 当年生叶; CR, 粗根; CS, 当年生茎; FR, 细根; ON, 一年生叶; OS, 一年生茎。CK, 对照, 不添加氮肥; N, 添加氮肥处理。图中不同小写字母表示同一器官内不同处理间差异显著(p < 0.05)。

Fig. 3 Effect of nitrogen addition on nitrogen (N) concentration (A), phosphorus (P) concentration (B), the pool sizes of N (C) and the pool sizes of P (D) in different organs (O) of Cunninghamia lanceolate seedlings (mean ± SE, n = 5). CN, current-year needles; CR, coarse roots; CS, current-year stems; FR, fine roots; ON, one-year-old needles; OS, one-year-old stems. CK, control, no nitrogen fertilizer addition; N, nitrogen addition. The different lowercase letters of the same organs indicate significant difference between treatments (p < 0.05).

图4 氮添加对杉木幼苗总生物量(A)、非结构性碳水化合物(NSC)(B)和结构性碳水化合物(SC)(C)库的地下与地上比值的影响(平均值±标准误, n = 5)。CK, 对照, 不添加氮肥; N, 添加氮肥处理。图中不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 4 Effect of nitrogen addition on the ratio of below- to aboveground total biomass (A), non-structural carbohydrates (NSC) pool size (B) and structural carbohydrates (SC) pool size (C) of Cunninghamia lanceolate seedlings (mean ± SE, n = 5). CK, control, no nitrogen fertilizer addition; N, nitrogen addition. Different lowercase letters indicate significant difference between treatments (p < 0.05).

表2 氮添加对杉木幼苗地上和地下部分生物量、非结构性碳水化合物(NSC)和结构性碳水化合物(SC)、氮(N)库和磷(P)库的影响

Table 2 Effect of nitrogen addition on the pool sizes of biomass, non-structural carbohydrates (NSC) and structural carbohydrates (SC)), nitrogen (N) and phosphorus (P) in the below- and aboveground of Cunninghamia lanceolate seedlings

器官 Organ	生物量 Biomass (g)	C (g)		N (g)	P (g)
器官 Organ	生物量 Biomass (g)	NSC	SC	N (g)	P (g)
CN	0	0	0	↑	↑
ON	0	0	0	0	0
CS	0	0	0	↑	0
OS	0	0	0	↑	↑
CR	0	0	0	0	0
FR	0	0	0	↑	0
地下总量(库) Belowground (pool)	0	0	0	↑	0
地上总量(库) Aboveground (pool)	0	0	0	↑	↑
地下/地上(库) Belowground/Aboveground (pool)	↓	0	↓	0	↓

图5 氮添加对杉木幼苗氮(N)库(A)和磷(P)库(B)的地下与地上比值的影响(平均值±标准误, n = 5)。CK, 对照, 不添加氮肥; N, 添加氮肥处理。图中不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 5 Effect of nitrogen addition on the ratio of below- to aboveground nitrogen (N) pool size (A) and phosphorus (P) pool size (B) of Cunninghamia lanceolate seedlings (mean ± SE, n = 5). CK, control, no nitrogen fertilizer addition; N, nitrogen addition. Different lowercase letters indicate significant difference between treatments (p < 0.05).

表3 氮添加对土壤pH、有效磷、铵态氮和硝态氮含量的影响(平均值±标准误, n = 5)

Table 3 Effect of nitrogen addition on soil pH, available phosphorus (AP), ammonium (NH4+-N) and nitrate nitrogen (NO3--N) concentrations of soil (mean ± SE, n = 5)

处理 Treatment	pH	有效磷 AP (mg·kg^-1)	铵态氮 NH₄⁺-N (mg·kg^-1)	硝态氮 NO₃^--N (mg·kg^-1)
CK	4.94 ± 0.09^a	0.28 ± 0.01^b	15.24 ± 0.49^b	4.92 ± 1.61^b
N	4.02 ± 0.04^b	0.40 ± 0.02^a	124.51 ± 7.42^a	61.75 ± 1.36^a

图6 氮添加对土壤脲酶(A)、亮氨酸氨基肽酶(B)、酸性磷酸酶(C)和N-乙酰氨基葡糖苷酶(D)活性的影响(平均值±标准误, n = 5)。CK, 对照, 不添加氮肥; N, 添加氮肥处理。图中不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 6 Effect of nitrogen addition on urease (A), Leucine aminopeptidase (B), acid phosphatase (C) and N-acetyl-glucosidase (NAG)(D) activity of soil (mean ± SE, n = 5). CK, control, no nitrogen fertilizer addition; N, nitrogen addition. Different lowercase letters indicate significant difference between treatments (p < 0.05).

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杉木幼苗生物量分配格局对氮添加的响应

Responses of biomass allocation patterns to nitrogen addition of Cunninghamia lanceolata seedlings

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器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

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器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0

器官 Organ	NSC (mg·g^-1)	可溶性糖 Soluble sugar (mg·g^-1)	淀粉 Starch (mg·g^-1)	N (mg·g^-1)	P (mg·g^-1)
CN	↓	↓	↓	↑	↑
ON	↓	↓	↓	↑	↑
CS	0	0	0	↑	0
OS	0	0	↓	↑	↑
CR	0	0	0	↑	0
FR	0	0	0	↑	0