氮磷添加对黑果枸杞（Lycium ruthenicum）营养器官非结构性碳水化合物特征的影响

doi:10.7522/j.issn.1000-694X.2020.00119

摘要/Abstract

摘要：

非结构性碳水化合物（NSC）在植物生长、繁殖、防御和生存等方面发挥着基础性作用。为探究黑果枸杞（Lycium ruthenicum）NSC积累的器官间规律及其对土壤氮磷供应水平的响应，以3年生黑果枸杞为对象，测定了3个氮磷比例和供应量条件下根茎叶NSC含量。结果表明：（1）黑果枸杞根、茎、叶NSC组成均以可溶性糖为主，淀粉约占37.3%；器官NSC含量粗根和细根>叶片>新梢>一二年生枝条，各器官可溶性糖含量粗根和细根>叶片>新梢>一年生枝>二年生枝；淀粉含量细根>粗根>二年生枝>一年生枝>叶片和新梢；糖淀比叶片>新梢>粗根>一年生枝>二年生枝>细根。（2）氮磷比和供应量对黑果枸杞根系可溶性糖含量无显著影响，对茎叶NSC含量的影响存在显著交互效应。氮磷比5∶1时，茎叶NSC含量随氮磷供应量升高而增加；氮磷比15∶1时，茎叶NSC含量随氮磷供应量增加显著降低；氮磷比45∶1时，茎叶NSC含量随供应量的增加而显著升高。（3）一年生枝NSC含量于萌芽前达到最大值（5.05% DW），于春梢旺盛生长期降至最低值（3.79% DW）；二年生枝NSC含量在萌芽前最低（4.23% DW），于秋季落叶期达到最大值（4.33% DW）。（4）粗根和叶片NSC含量与一二年生枝NSC含量负相关；新梢NSC含量与叶片NSC、二年生枝淀粉含量正相关；一年生枝可溶性糖含量与二年生枝可溶性糖含量负相关；二年生枝淀粉含量与根系淀粉和NSC含量负相关。黑果枸杞通过调整各器官NSC积累及分配应对生长季各器官能量需求和环境氮磷供应变化，在氮磷限制环境中将更多NSC储存在根系，可平衡当前碳同化与未来能量需求的矛盾，增强个体生存能力，也为黑果枸杞应对非生物逆境胁迫提供更大的灵活性。

关键词: 氮磷添加, 氮磷比, 非结构性碳水化合物, 黑果枸杞（Lycium ruthenicum）

Abstract:

Non-structural carbohydrates （NSC） played a fundamental role in plant growth， reproduction， defense and survival. To explore the effects of N and P addition on NSC of Lycium ruthenicum，a field manipulation experiment combined three nutrient supply levels （Low， Medium， High） and three N：P supply ratios （5∶1， 15∶1， 45∶1） was conducted on three-year-old L. ruthenicum. The results indicated that：（1） Soluble sugar was the major contributor to the total NSC in different organs， starch accounts for about 37.3%. On average， the concentrations of NSC in different organs followed the order of： coarse roots & fine roots>leaves>current-year>one-year& two-year branches. The concentrations of soluble sugar， starch and the ratio of soluble sugar to starch followed the order of coarse roots & fine roots>leaves>current-year branches>one-year branches>two-year branches； fine roots>coarse roots>two-year branches>one-year branches>leaves & current-year branches； leaves>current-year branches>coarse>one-year branches>two-year branches>fine roots respectively. （2） Changes in soluble sugar，starch and NSC concentrations in different organs of L. ruthenicum. reflected significant interactions between N∶P supply ratios and supply levels. When the N∶P supply ratios was 5∶1， the NSC concentrations of leaves and branches was higher under high supply level， when the N∶P was 15∶1， the NSC and its components concentrations of leaves，current-year and one-year branches in the low supply level is significantly higher than the high supply level， when the N∶P was 45∶1， the NSC concentrations of leaves，one-year and two-year branches in high level of supply is significantly higher than low supply level. N∶P supply ratios and supply level had no significant effect on root soluble sugar concentrations. （3） The NSC concentrations of one-year branches reached the maximum value （5.05% DW） before germination and dropped to the lowest value （3.79% DW） during the shoot growth period； NSC concentrations of two-year branches decreased before germination （4.23% DW） and reached the maximum value in the foliage period （4.33% DW）. （4） The NSC concentrations of coarse roots was negatively correlated with NSC concentrations of leaves、one and two-year branches； NSC concentrations of current-year branches was positively correlated with NSC concentrations of leaves and starch content of two-year branches； soluble sugar concentrations of one-year branches was negatively correlated with that of two-year branches； starch concentrations of two-year branches was negatively correlated with starch and NSC concentrations of coarse roots.When compared the impact of N and P addition on NSC concentrations of vegetative organs of L. ruthenicum， the results showed obvious organ difference adapted to N and P supply condition by adjusting accumulate and distribute NSC in different organs. Choosing to store more NSC in root system can enhance carbon balance and survivability in a nutrient-restricted environment， and also provide greater flexibility for the interaction with the non-biological environment.

Key words: nitrogen and phosphorus addition, N∶P, non-structural carbohydrate, Lycium ruthenicum

中图分类号:

Q948.1

李金霞, 朱亚男, 孙小妹, 韩国君, 陈年来. 氮磷添加对黑果枸杞（Lycium ruthenicum）营养器官非结构性碳水化合物特征的影响[J]. 中国沙漠, 2021, 41(2): 200-211.

Jinxia Li, Yanan Zhu, Xiaomei Sun, Guojun Han, Nianlai Chen. Effects of nitrogen and phosphorus addition on non-structural carbohydrate properties of vegetative organs in Lycium ruthenicum[J]. Journal of Desert Research, 2021, 41(2): 200-211.

图/表 9

图1 非结构性碳水化合物在黑果枸杞营养器官中的分布格局不同小写字母表示同一氮磷供应量或同一氮磷比不同器官间差异显著(P<0.05)

Fig.1 Patterns of NSC concentrations in vegetative organs of Lycium ruthenicum

图2 氮磷添加对黑果枸杞叶片非结构性碳水化合物特征的影响不同小写字母表示同一氮磷比不同供应量水平间差异显著，不同大写字母表示同一供应量下不同氮磷比间差异显著(P<0.05)

Fig.2 Effects of nitrogen and phosphorus addition on non-structural carbohydrates in leaves of Lycium ruthenicum

表1 氮磷比和供应量对黑果枸杞叶片非结构性碳水化合物的双因素方差分析

Table 1 Two-way ANOVA on the effects of N∶P supply ratio and overall supply level on non-structural carbohydrates in leaves of Lycium ruthenicum(F value)

因素	叶片
因素	可溶性糖	淀粉	NSC	糖/淀粉
供应比	28.2^***	5.8^*	14.1^***	3.5
供应量	1.9^**	0.6	1.4	1.4
供应比×供应量	14.1^***	5.7^*	9.9^***	4.4^*

图3 黑果枸杞一二年生枝条NSC含量的季节动态

Fig.3 Chronological changes of NSC and its components in the one-year and two-year branch of Lycium ruthenicum

表2 氮磷比和供应量对黑果枸杞枝条非结构性碳水化合物(NSC)的双因素方差分析

Table 2 Two-way ANOVA on the effects of N∶P supply ratio and overall supply level on non-structural carbohydrates in branches of Lycium ruthenicum(F value)

因素	新梢				一年生枝				二年生枝
因素	可溶性糖	淀粉	NSC	糖/淀粉	可溶性糖	淀粉	NSC	糖/淀粉	可溶性糖	淀粉	NSC	糖/淀粉
供应比（SR）	13.2^***	8.4^**	14.5^***	7.1^**	7.6^**	12.7^***	14.1^***	0.4	21.0^***	0.2	7.6^**	10.5^**
供应量（SL）	20.3^***	1.2	2.0	1.8	27.3^***	60.4^***	8.2^**	104.4^***	12.4^***	48.0^***	8.8^**	94.0^***
供应比×供应量	26.4^***	8.2^**	14.6^***	11.5^***	12.6^***	8.6^***	12.8^***	5.2^**	3.9^*	10.6^***	5.2^**	8.9^***

图4 氮磷添加对黑果枸杞不同年龄枝条非结构性碳水化合物特征的影响不同小写字母表示同一氮磷比不同供应量水平间差异显著，不同大写字母表示同一供应量下不同氮磷比间差异显著(P<0.05)

Fig.4 Effects of nitrogen and phosphorus addition on non-structural carbohydrates in different age branches of Lycium ruthenicum

表3 氮磷比和供应量对黑果枸杞根系非结构性碳水化合物(NSC)的双因素方差分析

Table 3 Two-way ANOVA on the effects of N∶P supply ratio and overall supply level on non-structural carbohydrates in roots of Lycium ruthenicum(F value)

因素	细根				粗根
因素	可溶性糖	淀粉	NSC	糖/淀粉	可溶性糖	淀粉	NSC	糖/淀粉
供应比	1.2	16.0^***	6.2^**	25.5^***	2.6	6.7^**	4.5^*	10.7^**
供应量	0.8	33.3^***	13.5^***	51.5^***	1.4	25.8^***	22.2^***	35.6^***
供应比×供应量	0.7	6.7^**	3.9^*	9.0^***	0.3	9.6^***	10.0^***	11.9^***

图5 氮磷添加对黑果枸杞粗根和细根非结构性碳水化合物特征的影响不同小写字母表示同一氮磷比不同供应量水平间差异显著，不同大写字母表示同一供应量下不同氮磷比间差异显著(P<0.05)

Fig.5 Effects of nitrogen and phosphorus addition on non-structural carbohydrates in coarse and fine roots of Lycium ruthenicum

图6 土壤化学性质与不同器官中NSC及其组分含量、可溶性糖/淀粉的RDA排序1、2、3、4、5、6分别代表叶片、新梢、一年生枝、二年生枝、粗根、细根；SS、ST、NSC、SS/ST分别代表可溶性糖、淀粉、非结构性碳水化合物、可溶性糖/淀粉

Fig.6 RDA sort graph of soil chemical properties and NSC and its components and soluble sugar/starch in different organs

参考文献 44

1	Dietze M C，Sala A，Carbone M S，et al.Nonstructural carbon in woody plants［J］.Annual Review of Plant Biology，2014，65（1）： 667-687.
2	刘青青，马祥庆，黄智军，等.光强对杉木幼苗形态特征和叶片非结构性碳含量的影响［J］.生态学报，2019，39（12）：4455-4462.
3	Chapin F S，Schulze E D，Mooney H A.The ecology and economics of storage in plants［J］.Annual Review of Ecology，Evolution，and Systematics，1990，21（1）：423-447.
4	Hartmann H，Trumbore S E.Understanding the roles of nonstructural carbohydrates in forest trees-from what we can measure to what we want to know［J］.New Phytologist，2016，211（2）：386-403.
5	章异平，曹鹏鹤，徐军亮，等.秦岭东段栓皮栎叶片非结构性碳水化合物含量的季节动态［J］.生态学报，2019，39（19）：7274-7282.
6	Gusewell S.N∶P ratios in terrestrial plants： variation and functional significance［J］.New Phytologist，2004，164（2）： 243-266.
7	林婉奇，蔡金桓，薛立.氮磷添加与不同栽植密度交互对樟树幼苗土壤化学性质的短期影响［J］.生态学报，2019，39（24）：9162-9170.
8	Goodsman D W，Lieffers V J，Landhäusser S M，et al.Fertilization of lodgepole pine trees increased diameter growth but reduced root carbohydrate concentrations［J］.Forest ecology and management，2010，260（10）：1914-1920.
9	史顺增，熊德成，冯建新，等.模拟氮沉降对杉木幼苗细根的生理生态影响［J］.生态学报，2017，37（1）：74-83.
10	Kobe R K，Iyer M，Walters M B.Optimal partitioning theory revisited：nonstructural carbohydrates dominate root mass responses to nitrogen［J］.Ecology，2010，91（1）：166-179.
11	Nanamori M，Shinano T，Wasaki J，et al.Low phosphorus tolerance mechanisms ： phosphorus recycling and photosynthate partitioning in the tropical forage grass，brachiaria hybrid cultivar mulato compared with rice［J］.Plant and Cell Physiology，2004，45（4）：460-469.
12	Woodruff D R，Meinzer F C.Water stress，shoot growth and storage of non-structural carbohydrates along a tree height gradient in a tall conifer［J］.Plant，Cell & Environment，2011，34（11）：1920-1930.
13	Guo Y Y，Nie H S，Yu H Y，et al.Effect of salt stress on the growth and photosystem II photochemical characteristics of Lycium ruthenicum Murr.seedlings［J］.Photosynthetica，2019，57（2）：564-571.
14	Güsewell S.High nitrogen：phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges［J］.New Phytologist，2005，166（2）：537-550.
15	Li M，Xiao W，Wang S，et al.Mobile carbohydrates in Himalayan treeline trees I.evidence for carbon gain limitation but not for growth limitation［J］.Tree Physiology，2008，28（8）：1287-1296.
16	鲍士旦.土壤农化分析［M］.北京：中国农业出版社，2007：268-270，389-391.
17	Cannell M G，Dewar R C.Carbon allocation in trees：a review of concepts for modelling［J］.Advances in Ecological Research，1994：59-104.
18	Wurth M K，Pelaezriedl S，Wright S J，et al.Non-structural carbohydrate pools in a tropical forest［J］.Oecologia，2005，143（1）：11-24.
19	Boldingh H，Smith G S，Klages K.Seasonal concentrations of non-structural carbohydrates of five actinidia species in fruit，leaf and fine root tissue［J］.Annals of Botany，2000，85（4）：469-476.
20	Sala A，Mencuccini M.Ecosystem science：plump trees win under drought［J］.Nature Climate Change，2014，4（8）：666-667.
21	Lacointe A，Kajji A，Daudet F A，et al.Mobilization of carbon reserves in young walnut trees［J］.Acta Botanica Gallica，1993，140（4）：435-441.
22	王凯，雷虹，夏扬，等.杨树幼苗非结构性碳水化合物对增加降水和氮添加的响应［J］.应用生态学报，2017，28（2）：399-407.
23	Schadel C，Blochl A，Richter A，et al.Short-term dynamics of nonstructural carbohydrates and hemicelluloses in young branches of temperate forest trees during bud break［J］.Tree Physiology，2009，29（7）：901-911.
24	Gaucher C，Gougeon S，Mauffette Y，et al.Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple （Acer saccharum） and yellow birch （Betula alleghaniensis） seedlings［J］.Tree Physiology，2005，25（1）：93-100.
25	Kozlowski T T.Carbohydrate sources and sinks in woody plants ［J］.Botanical Review，1992，58（2）：107-222.
26	Eissenstat D M，Yanai R D.The Ecology of Root Lifespan ［J］.Advances in Ecological Research，1997，27：1-60.
27	Rothstein D E，Zak D R，Pregitzer K S，et al.Kinetics of nitrogen uptake by populus tremuloides in relation to atmospheric CO₂ and soil nitrogen availability［J］.Tree Physiology，2000，20（4）：265-270.
28	Limin Y U，Wang C，Wang X，et al.Allocation of nonstructural carbohydrates for three temperate tree species in Northeast China：allocation of nonstructural carbohydrates for three temperate tree species in Northeast China［J］.Chinese Journal of Plant Ecology，2011，35（12）：1245-1255.
29	Kobe R K，Iyer M，Walters M B.Optimal partitioning theory revisited：nonstructural carbohydrates dominate root mass responses to nitrogen［J］.Ecology，2010，91（1）：166-179.
30	Klein T，Vitasse Y，Hoch G.Coordination between growth，phenology and carbon storage in three coexisting deciduous tree species in a temperate forest［J］.Tree Physiology，2016，36（7）：847-855.
31	张海燕，王传宽，王兴昌.温带12个树种新老树枝非结构性碳水化合物浓度比较［J］.生态学报，2013，33（18）：5675-5685.
32	Keel S G，Schadel C.Expanding leaves of mature deciduous forest trees rapidly become autotrophic［J］.Tree Physiology，2010，30（10）：1253-1259.
33	章异平，海旭莹，徐军亮，等.秦岭东段栓皮栎枝条非结构性碳水化合物含量的季节动态［J］.植物生态学报，2019，43（6）：521-531.
34	Verdaguer D.Root starch storage and allocation patterns in seeder and resprouter seedlings of two Cape Erica （Ericaceae） species［J］.American Journal of Botany，2002，89（8）：1189-1196.
35	Guo Q X，Li J Y，Zhang Y X，et al.Species-specific competition and N fertilization regulate nonstructural carbohydrate contents in two Larix species［J］.Forest Ecology and Management，2016，364：60-69.
36	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.
37	王雪，雒文涛，庾强，等.半干旱典型草原养分添加对优势物种叶片氮磷及非结构性碳水化合物含量的影响［J］.生态学杂志，2014，33（7）：1795-1802.
38	克热木·伊力，新居直.不同氮素施用量对桃树叶碳水化合物含量积累的影响［J］.新疆农业大学学报，2000（1）：37-41.
39	史顺增，熊德成，邓飞，等.土壤增温、氮添加及其交互作用对杉木幼苗细根生产的影响［J］.植物生态学报，2017，41（2）：186-195.
40	Coley M L D.Benefits of the carbon-nutrient balance hypothesis ［J］.Oikos，2002，98（3）：534-536.
41	Gebbing T，Schnyder H.Pre-anthesis reserve utilization for protein and carbohydrate synthesis in grains of wheat［J］.Plant Physiology，1999，121（3）：871-878.
42	Peng Z T，Chen M X，Huang Z J，et al.Non-structural carbohydrates regulated by nitrogen and phosphorus fertilization varied with organs and fertilizer levels in Moringa oleifera seedlings［J］.Journal of Plant Growth Regulation，2020：1-10.
43	Woodruff D R，Meinzer F C，Marias D E，et al.Linking nonstructural carbohydrate dynamics to gas exchange and leaf hydraulic behavior in Pinus edulis and Juniperus monosperma［J］.New Phytologist，2015，206（1）：411-421.
44	White A E，Li Y J，Griskevicius V，et al.Putting all your eggs in one basket： life-history strategies，bet hedging，and diversification［J］.Psychological Science，2013，24（5）：715-722.

[1]	赵泽芳, 卫海燕, 郭彦龙, 赵泽斌, 庞国锦, 马媛, 顾蔚. 黑果枸杞(Lycium ruthenicum)分布对气候变化的响应及其种植适宜性[J]. 中国沙漠, 2017, 37(5): 902-909.
[2]	罗君, 彭飞, 王涛, 薛娴, 黄翠华. 黑果枸杞(Lycium ruthenicum)种子萌发及幼苗生长对盐胁迫的响应[J]. 中国沙漠, 2017, 37(2): 261-267.
[3]	朱秉坚, 尹本丰, 张元明. 不同微生境下齿肋赤藓(Syntrichia caninervis)非结构性碳水化合物含量的季节动态[J]. 中国沙漠, 2017, 37(2): 268-275.
[4]	强生斌, 周瑞莲, 梁慧敏. 海岸植物滨麦(Leymus mollis)和肾叶打碗花(Calystegia soldanella)根叶碳水化合物含量与土壤Na⁺含量的关系[J]. 中国沙漠, 2016, 36(6): 1613-1621.
[5]	邬畏;何兴东*;周启星. 生态系统氮磷比化学计量特征研究进展[J]. 中国沙漠, 2010, 30(2): 296-302.