霸王根茎叶非结构性碳与C:N:P计量特征对干旱的响应

doi:10.12118/j.issn.1000–6060.2021.01.25

干旱区地理 ›› 2021, Vol. 44 ›› Issue (1): 240-249.doi: 10.12118/j.issn.1000–6060.2021.01.25

霸王根茎叶非结构性碳与C:N:P计量特征对干旱的响应

孙小妹¹(),何明珠²,周彬³,李金霞¹,陈年来¹()

1.甘肃农业大学资源与环境学院,甘肃兰州 730070
2.中国科学院西北生态环境资源研究院沙坡头沙漠研究试验站,甘肃兰州 730000
3.山东农业大学资源与环境学院,山东泰安 271018

收稿日期:2020-02-27 修回日期:2020-06-28 出版日期:2021-01-25 发布日期:2021-03-09
通讯作者: 陈年来
作者简介:孙小妹（1986-）,女,博士,讲师,研究方向为恢复生态学. E-mail: sunxm@gsau.edu.cn
基金资助:
甘肃农业大学科技创新基金(学科建设专项基金)(GSAU-XKJS-2018-214/198);国家自然科学基金(41671103);横向项目(XZ 20191206)

Non-structural carbohydrates and C:N:P stoichiometry of roots, stems, and leaves of Zygophyllum xanthoxylon in responses to xeric condition

SUN Xiaomei¹(),HE Mingzhu²,ZHOU Bin³,LI Jinxia¹,CHEN Nianlai¹()

1. College of Resource and Environmental Sciences, Gansu Agricultural University, Lanzhou 730070, Gansu, China
2. Shapotou Desert Experiment and Research Station, Northwest Institute of Eco-Environment and Researches , Chinese Academy of Sciences, Lanzhou 730000, Gansu, China
3. College of Resources and Environment, Shandong Agricultural University, Tai’an 271018, Shandong, China

Received:2020-02-27 Revised:2020-06-28 Online:2021-01-25 Published:2021-03-09
Contact: Nianlai CHEN

摘要/Abstract

摘要：

通过选取阿拉善地区3个典型霸王群落为研究对象,研究霸王器官间（根、茎、叶）生态化学计量比特征及非结构性碳水化合物（Non-structural carbohydrates,NSCs）的变化,以便加深对霸王生存策略的理解,更好的服务于荒漠生态系统的生态恢复。结果表明：霸王茎和叶中可溶性糖（Soluble sugar,SS）、淀粉（Starch,ST）和NSCs含量随干旱加剧均显著降低。根系中SS和NSCs在3个样点间差异不显著,ST先降低后增加,ST含量在Plot3中比在Plot1和Plot2中高19.3%和31.2%。干旱使霸王根、茎、叶中N、P含量显著下降,使根、茎、叶中C:N和C:P显著增高,叶片N:P<14。Plot1中霸王器官间SS、SS:ST和NSCs含量表现为叶片>根>茎,ST含量为茎>根>叶片;Plot2和Plot3中霸王器官间SS、ST和NSCs含量均为根>茎>叶片。3个样点中叶片相较于茎干和根系具有高的N、P含量和低的C:N和C:P。NSCs与C:N:P间的关系表明：根、茎、叶中的N含量与SS正相关,叶片N含量与根系ST负相关。以上结果表明霸王的茎干和根系可能扮演“营养库”的角色,霸王通过提高N、P利用效率来减缓水分亏缺使其生长受到N限制的影响。霸王通过调节器官间NSCs的积累及分配来适应干旱,采取将根系中ST转化为SS来调节渗透势的策略;并随干旱加剧将更多的碳水化合物投资分配到根部,以ST的形式存储,而N是影响霸王根系中ST与SS间转化和ST存储的关键元素。

关键词: 非结构性碳水化合物, 生态化学计量比特征, 器官关联性, 干旱适应策略

Abstract:

In arid ecosystems, high temperatures and low rainfall result in reduced carbon fixation. Non-structural carbohydrates (NSCs) (including soluble sugars and starch) represent the products of plant photosynthesis and are mainly involved in the balance between C acquisition and expenditure in life processes. NSCs also reflect the amount of plant carbohydrates that can be used to resist adverse environmental conditions. The C:N:P stoichiometry of plants is associated with important ecological processes, such as an organism’s ability to adapt to environmental stresses. Leaf N is the basis of chlorophyll formation for direct use in photosynthesis. Plant P is indispensable for the transportation of photosynthetic products. Thus, the C:N:P variations in plants are directly affected by the rate of photosynthesis. Leaf N concentration has been found to be positively correlated with NSC fixation ability, and P has been identified as the key element of plant metabolism. Accordingly, leaf photosynthetic capacity and NSC synthesis are not only affected by leaf N concentration but also closely related to P concentration. Therefore, it is important to study the relationship between NSCs and C:N:P stoichiometry in Zygophyllum xanthoxylon to improve our understanding of its survival and growth strategies to xeric conditions. Samples were collected from three dominant communities of Z. xanthoxylon across Ningxia and Inner Mongolia Provinces in northwestern China that differed in mean annual relative air moisture (Plot 1>Plot 2>Plot 3). Plant and soil samples were collected during the growing season (August) of 2014. Soluble sugars and starch are commonly studied NSCs. Plant samples were divided into leaves, stems, and roots to analyze the concentrations of starch (ST), soluble sugar (SS), C, N, and P. The concentrations of SS, ST, and NSCs in stems and leaves decreased from Plot 1 to Plot 3. N and P concentrations decreased, whereas C:N and C:P ratios increased with increasing xeric conditions in roots, stems, and leaves. N:P ratios in leaves were all below 13, lower than the critical ratio of 14 in all three plots, suggesting that N is the limiting factor for the growth of Z. xanthoxylon in xeric environments. Variations were inconsistent in NSCs and C:N:P stoichiometry among the organs. N and P concentrations in leaves were higher than those in stems and roots, while C:N and C:P ratios in leaves were lower than those in stems and roots in three plots. The correlation of C:N:P stoichiometry with NSCs showed that N concentrations positively correlated with SS concentrations in roots, stems, and leaves, whereas N concentrations in leaves negatively correlated with ST in roots. Stems and roots may act as nutrient sinks, and Z. xanthoxylon may alleviate water deficit-caused N limitation by increasing N and P use efficiency. Z. xanthoxylon adapted to drought by regulating the accumulation and distribution of NSCs among organs. Osmotic potential is regulated by conversions between ST and SS in the root system; more carbohydrates were allocated to roots and stored as ST with enhancing drought stress. N is the key factor in the transformation of ST to SS and ST storage in the root system.

Key words: non-structural carbohydrates, C:N:P ratios, organ relevance, adaptive strategy of xeric condition

孙小妹,何明珠,周彬,李金霞,陈年来. 霸王根茎叶非结构性碳与C:N:P计量特征对干旱的响应[J]. 干旱区地理, 2021, 44(1): 240-249.

SUN Xiaomei,HE Mingzhu,ZHOU Bin,LI Jinxia,CHEN Nianlai. Non-structural carbohydrates and C:N:P stoichiometry of roots, stems, and leaves of Zygophyllum xanthoxylon in responses to xeric condition[J]. Arid Land Geography, 2021, 44(1): 240-249.

图/表 7

图1

表1

表2

图2

图3

表3

图4

参考文献 44

[1]	Choat B, Jansen S, Brodribb T J, et al. Global convergence in the vulnerability of forests to drought[J]. Nature, 2012,491(7426):752-755. doi: 10.1038/nature11688 pmid: 23172141
[2]	Clark J S, Bell D M, Kwit M, et al. Individual-scale inference to anticipate climate-change vulnerability of biodiversity[J]. Philosophical Transactions of the Royal Society B Biological Sciences, 2012,367(1586):236.
[3]	Stitt M, Schulze D. Does rubisco control the rate of photosynjournal and plant growth? An exercise in molecular ecophysiology[J]. Plant Cell Environment , 1994,17(5):465-487.
[4]	Warren C R, Adams M A, Chen Z L. Is photosynjournal related to concentrations of nitrogen and rubisco in leaves of Australian native plants?[J]. Functional Plant Biology, 2000,27(5):407-416.
[5]	Lambers H, Chapin F S, Pons T L. Plant physiological ecology[M]. New York: Springer, 1998: 185.
[6]	Azcon R, Gomez M, Tobar R. Effects of nitrogen source on growth, nutrition, photosynthetic rate and nitrogen metabolism of mycorrhizal and phosphorus-fertilized plants of Lactuca sativa L.[J]. New Phytologist, 1992,121(2):227-234.
[7]	李从娟, 徐新文, 孙永强, 等. 不同生境下三种荒漠植物叶片及土壤C、N、P的化学计量特征[J]. 干旱区地理, 2014,37(5):996-1004.
	[ Li Congjuan, Xu Xinwen, Sun Yongqiang, et al. Stoichiometric characteristics of C, N, P for three desert plants leaf and soil at different habitats[J]. Arid Land Geography, 2014, 37(5): 996-1004].
[8]	周欣, 左小安, 赵学勇, 等. 科尔沁沙地不同生境植物及叶片的C、N元素计量特征[J]. 干旱区地理, 2015,38(3):565-575.
	[ Zhou Xin, Zuo Xiao’an, Zhao Xueyong, et al. Ecological stoichiometry of plant and leaf carbon and nitrogen in different habitats of Horqin Sandy Land[J]. Arid Land Geography, 2015, 38(3): 565-575].
[9]	Leuzinger S, Bigler C, Wolf A, et al. Poor methodology for predicting large-scale tree die-off[J]. Proceedings of the National Academy of Sciences of the United States of America, 2009,106(38):106-107.
[10]	Zhang T, Cao Y, Chen Y, et al. Non-structural carbohydrate dynamics in Robinia pseudoacacia, saplings under three levels of continuous drought stress[J]. Trees, 2015,29(6):1837-1849.
[11]	Arx G V, Arzac A, Fonti P, et al. Responses of sapwood ray parenchyma and non-structural carbohydrates of Pinus sylvestris to drought and long-term irrigation[J]. Functional Ecology, 2017,31(7):1371-1382.
[12]	Xie H, Yu M, Cheng X. Leaf non-structural carbohydrate allocation and C:N:P stoichiometry in response to light acclimation in seedlings of two subtropical shade-tolerant tree species[J]. Plant Physiology and Biochemistry, 2018,124(1):146-154.
[13]	Sala A, Woodruff D R, Meinzer F C. Carbon dynamics in trees: Feast or famine[J]. Tree Physiology, 2012,32(6):764-775. doi: 10.1093/treephys/tpr143 pmid: 22302370
[14]	Reich P B, Oleksyn J. Global patterns of plant leaf N and P in relation to temperature and latitude[J]. Proceedings of the National Academy of Sciences. 2004,101(30):11001-11006. doi: 10.1073/pnas.0403588101
[15]	Mcgroddy M E, Daufresne T, Hedin L O. Scaling of C: N: P stoichiometry in forests worldwide: Implications of terrestrial Redfield-type ratios[J]. Ecology, 2004,85(5):2390-2401. doi: 10.1890/03-0351
[16]	Chapin F S. The mineral nutrition of wild plants[J]. Annual Review Ecology and Systematics. 1980,11(1):233-260. doi: 10.1146/annurev.es.11.110180.001313
[17]	Chapin F S, Schulze E D, Mooney H A. The ecology and economics of storage in plants[J]. Annual Review Ecology and Systematics. 1990,21(1):423-447.
[18]	Li F, Hu J, Xie Y, et al. Foliar stoichiometry of carbon, nitrogen, and phosphorus in wetland sedge Carex brevicuspis along a small-scale elevation gradient[J]. Ecological Indicators, 2018,92(9):322-329.
[19]	Lu X T, Freschet G T, Kazakou E, et al. Contrasting responses in leaf nutrient-use strategies of two dominant grass species along a 30-yr temperate steppe grazing exclusion chronosequence[J]. Plant and Soil, 2015,387(2):69-79.
[20]	Sala A, Hoch G. Height-related growth declines in Ponderosa pine are not due to carbon limitation[J]. Plant Cell and Environment, 2010,32(1):22-30.
[21]	He M Z, Dijkstra F A, Zhang K, et al. Influence of life form, taxonomy, climate, and soil properties on shoot and root concentrations of 11 elements in herbaceous plants in a temperate desert[J]. Plant and Soil, 2016,398(1):339-350.
[22]	He M Z, Zhang K, Tan H, et al. Nutrient levels within leaves, stems, and roots of the xeric species Reaumuria soongorica in relation to geographical, climatic, and soil conditions[J]. Ecology and Evolution, 2015,5(7):1494-1503. doi: 10.1002/ece3.1441 pmid: 25897388
[23]	李培广, 周海燕, 陈翠云, 等. 阿拉善荒漠优势植物可溶性糖的季节变化[J]. 生态学杂志, 2012,31(12):3018-3023.
	[ Li Peiguang, Zhou Haiyan, Chen Cuiyun, et al. Seasonal variations of soluble sugars in dominant plant species in Alxa Desert of northwest China[J]. Chinese Journal of Ecology, 2012,31(12):3018-3023. ]
[24]	牛得草, 陈鸿洋, 江世高, 等. 荒漠植物霸王(Zygophyllum xanthoxylum)不同大小叶片C、N、P化学计量特征[J]. 中国沙漠, 2013,33(3):703-709.
	[ Niu Decao, Chen Hongyang, Jiang Shigao, et al. Stoichiometric traits of C, N, P of leaves in desert shrub Zygophyllum xanthoxylum[J]. Journal of Desert Research, 2013,33(3):703-709. ]
[25]	牛得草, 李茜, 江世高, 等. 阿拉善荒漠区6种主要灌木植物叶片C: N: P化学计量比的季节变化[J]. 植物生态学报, 2013,37(4):317-325.
	[ Niu Decao, Li Xi, Jiang Shigao, et al. Seasonal variations of leaf C:N:P stoichiometry of six shrubs in desert of China’s Alxa Plateau[J]. Chinese Journal of Plant Ecology, 2013,37(4):317-325. ]
[26]	Ramirezbriones E, Rodriguzmacias R, Salcedoperez E, et al. Seasonal variation in non-structural carbohydrates, sucrolytic activity and secondary metabolites in deciduous and perennial Diospyros species sampled in Western Mexico[J]. PloS One, 2017,12(10):1-24.
[27]	O’brien M J, Leuzinger S, Philipson C D, et al. Drought survival of tropical tree seedlings enhanced by non-structural carbohydrate levels[J]. Nature Climate Change, 2014,4(8):710-714. doi: 10.1038/NCLIMATE2281
[28]	Sala A, Hoch G. Height-related growth declines in Ponderosa pine are not due to carbon limitation[J]. Plant Cell and Environment, 2010,32(1):22-30. doi: 10.1111/pce.2009.32.issue-1
[29]	Zhang T, Cao Y, Chen Y, et al. Non-structural carbohydrate dynamics in Robinia pseudoacacia, saplings under three levels of continuous drought stress[J]. Trees, 2015,29(6):1837-184.
[30]	王延平, 许坛, 朱婉芮, 等. 杨树细根碳、氮含量的季节动态及代际差异. 应用生态学报, 2015,26(11):21-29.
	[ Wang Yanping, Xu Tan, Zhu Wanru, et al. Seasonal dynamics of carbon and nitrogen in fine roots and their differences between successive rotation poplar plantations[J]. Chinese Journal of Plant Ecology, 2015,26(11):21-29]
[31]	郑云普, 王贺新, 娄鑫, 等. 木本植物非结构性碳水化合物变化及其影响因子研究进展. 应用生态学报, 2014,25(4):1188-1196.
	[ Zheng Yunpu, Wang Hexin, Lou Xin, et al. Changes of non-structural carbohydrates and its impact factors in trees: A review[J]. Chinese Journal of Plant Ecology, 2014,25(4):1188-1196]
[32]	Ai Z M, Xue S, Wang G L, et al. Responses of non-structural carbohydrates and C:N:P stoichiometry of Bothriochloa ischaemum to nitrogen addition on the Loess Plateau, China[J]. Journal of Plant Growth Regulation, 2017,36(3):714-722. doi: 10.1007/s00344-017-9673-y
[33]	董蕾, 李吉跃. 植物干旱胁迫下水分代谢、碳饥饿与死亡机理. 生态学报, 2013,33(18):5477-5483. doi: 10.5846/stxb201304270839
	[ Dong Lei, Li Jiyue. Relationship among drought, hydraulic metabolic, carbon starvation and vegetation mortality[J]. Acta Ecologica Sinica, 2013,33(18):5477-5483] doi: 10.5846/stxb201304270839
[34]	Drenovsky R E, James J J, Richards J H. Variation in nutrient resorption by desert shrubs[J]. Journal of Arid Environment, 2010,74(11):1564-1568.
[35]	He M Z, Dijkstra F A. Drought effect on plant nitrogen and phosphorus: A meta-analysis[J]. New Phytologist, 2014,204(4):924-931.
[36]	Hartley A N, Bargerj B, Okin G. Dryland ecosystems [C]//Marschner P, Rengel Z. Nutrient Cycling in Terrestrial Ecosystems. Berlin & Heidelberg: Springer, 2007: 271-308.
[37]	Sterner R W, Elser J J. Ecological stoichiometry: The biology of elements from molecules to the biosphere[M]. Princeton: Princeton University Press, 2002: 357.
[38]	Vitousek P. Nutrient cycling and nutrient use efficiency[J]. American Naturalist, 1982,119(4):553-572.
[39]	王凯, 沈潮, 孙冰, 等. 干旱胁迫对科尔沁沙地榆树幼苗C、N、P化学计量特征的影响[J]. 应用生态学报. 2018,29(7):2286-2294.
	[ Wang Kai, Shen Chao, Sun Bin, et al. Effects of drought stress on C, N and P stoichiometry of Ulmus pumila seedlings in Horqin Sandy Land[J]. Chinese Journal of Applied Ecology, 2018,29(7):2286-2294 ]
[40]	Von A G, Archer S R, Hughes M K. Long-term functional plasticity in plant hydraulic architecture in response to supplemental moisture[J]. Annals of Botany, 2012,109(6):1091. doi: 10.1093/aob/mcs030 pmid: 22396436
[41]	于丽敏, 王传宽, 王兴昌. 三种温带树种非结构性碳水化合物的分配[J]. 植物生态学报, 2011,35(12):1245-1255.
	[ Yu Limin, Wang Chuankuan, Wang Xinchang. Allocation of nonstructural carbohydrates for three temperate tree species in northeast China[J]. Chinese Journal of Plant Ecology, 2011,35(12):1245-1255. ]
[42]	Farooq M, Hussain M, Wahid A, et al. Drought stress in plants: An overview[M]. Berlin & Heidelberg: Springer, 2012: 288.
[43]	Farooq M, Wahid A, Kobayashi N, et al. Plant drought stress: Effects, mechanisms and management[J]. Agronmy for Sustainable Development. 2009,29(1):185-212.
[44]	Thompson K E N, Parkinson J A, Band S R, et al. A comparative study of leaf nutrient concentrations in a regional herbaceous flora[J]. New Phytologist, 1997,136(4):679-689.

深度/cm	土壤有机质			土壤全氮			土壤全磷
深度/cm	Plot1	Plot2	Plot3	Plot1	Plot2	Plot3	Plot1	Plot2	Plot3
0~1	21.25±0.61a	20.56±0.20a	18.24±0.19b	0.14±0.005a	0.13±0.004ab	0.12±0.005b	0.73±0.007a	0.67±0.016a	0.44±0.027b
1~10	19.44±0.34a	17.89±0.21ab	16.81±0.74b	0.13±0.0005a	0.08±0.0009b	0.07±0.001c	0.59±0.004a	0.59±0.017a	0.39±0.015b
10~20	17.32±0.53a	15.34±0.22b	14.58±0.47b	0.12±0.003a	0.08±0.002b	0.06±0.003c	0.58±0.001a	0.57±0.004a	0.26±0.009b
20~40	16.15±0.71a	13.39±0.72b	12.87±0.07b	0.11±0.002a	0.07±0.002b	0.05±0.001c	0.57±0.002a	0.54±0.009b	0.19±0.003c
40~60	13.67±0.27a	11.41±0.08b	10.33±0.23c	0.09±0.002a	0.05±0.002b	0.05±0.0004c	0.52±0.007a	0.49±0.004b	0.17±0.005c
60~80		9.30±0.10a	8.87±0.05b		0.05±0.002a	0.04±0.001b		0.41±0.006a	0.17±0.007b
80~100			6.45±0.05			0.04±0.001			0.08±0.008

样点	植被类型	纬度/N	经度/E	海拔/m	年均温/℃	年均降雨量/mm	大于10 ℃的积温/℃	空气相对湿度/%
Plot1	霸王群落	39°53′2.04″	103°38′27.96″	1511	6.73	136.97	3149.47	26.40
Plot2	霸王群落	39°48′37.8″	105°3′57.96″	1477	6.91	117.57	3227.49	23.63
Plot3	霸王群落	39°56′40.56″	102°16′21.36″	1344	8.79	112.00	3749.43	22.73

器官		SS	ST	NSCs	C	N	P	C:N	C:P	N:P
根	SS	1.000
	ST	-0.222	1.000
	NSCs	0.675^*	0.57	1.000
	C	-0.577	0.818^**	0.132	1.000
	N	0.682^*	-0.734^*	0.019	-0.918^**	1.000
	P	0.59	-0.522	0.102	-0.677^*	0.893^**	1.000
	C:N	-0.63	0.786^*	0.064	0.908^**	-0.979^**	-0.832^**	1.000
	C:P	-0.651	0.722^*	-0.002	0.897^**	-0.994^*	-0.926^**	0.962^**	1.000
	N:P	0.477	-0.711^*	-0.136	-0.826^**	0.666^*	0.266	-0.745^*	-0.595	1.000
茎	SS	1.000
	ST	0.433	1.000
	NSCs	0.894^**	0.792^*	1.000
	C	0.01	-0.594	-0.289	1.000
	N	0.791^*	0.46	0.765^*	-0.045	1.000
	P	0.748^*	0.652^*	0.832^**	-0.272	0.955^**	1.000
	C:N	-0.763^*	-0.343	-0.688^*	-0.058	-0.987^**	-0.899^**	1.000
	C:P	-0.752^*	-0.467	-0.742^*	0.054	-0.995^**	-0.946^**	0.989^**	1.000
	N:P	0.248	-0.605	0.133	0.755^*	0.265	-0.032	-0.401	-0.270	1.000
叶	SS	1.000
	ST	0.941^**	1.000
	NSCs	0.997^**	0.965^**	1.000
	C	0.741^*	0.659^*	0.729^*	1.000
	N	0.819^**	0.761^*	0.814^**	0.966^**	1.000
	P	0.802^*	0.782^*	0.806^**	0.96^**	0.987^**	1.000
	C:N	-0.747^*	-0.679^*	-0.743^*	-0.932^**	-0.984^**	-0.960^**	1.000
	C:P	-0.749^*	-0.705^*	-0.746^*	-0.973^**	-0.971^**	-0.985^**	0.943^**	1.000
	N:P	-0.715^*	-0.688^*	-0.716^*	-0.936^**	-0.904^**	-0.946^**	0.847^**	0.975^**	1.000

霸王根茎叶非结构性碳与C:N:P计量特征对干旱的响应

Non-structural carbohydrates and C:N:P stoichiometry of roots, stems, and leaves of Zygophyllum xanthoxylon in responses to xeric condition

RichHTML

PDF (PC)

赞

可视化

摘要/Abstract

引用本文

使用本文

图/表 7

参考文献 44

相关文章 0

Metrics

本文评价

推荐阅读 0