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

doi:10.17521/cjpe.2018.0325

植物生态学报 ›› 2019, Vol. 43 ›› Issue (6): 521-531.DOI: 10.17521/cjpe.2018.0325

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

章异平¹,海旭莹^1,²,徐军亮^1,^*(),吴文霞¹,曹鹏鹤^1,³,安文静⁴

1 河南科技大学林学院, 河南洛阳 471023
2 西北农林科技大学林学院, 陕西杨凌 712100
3 南京林业大学生物与环境学院, 南京 210037
4 北京市京西林场, 北京 102300

收稿日期:2018-12-31 修回日期:2019-05-30 出版日期:2019-06-20 发布日期:2019-09-30
通讯作者: 徐军亮
作者简介:ORCID: 章异平: 0000-0001-5666-8198
基金资助:
国家自然科学基金(41401063);国家自然科学基金(41801026);国家留学基金(201808410575);国家留学基金(201908410061)

Seasonal dynamics of non-structural carbohydrate content in branch of Quercus variabilis growing in east Qinling Mountain range

ZHANG Yi-Ping¹,HAI Xu-Ying^1,²,XU Jun-Liang^1,^*(),WU Wen-Xia¹,CAO Peng-He^1,³,AN Wen-Jing⁴

1 College of Forestry, Henan University of Science and Technology, Luoyang, Henan 471023, China
2 College of Forestry, Northwest Agriculture and Forestry University, Yangling, Shaanxi 712100, China
3 College of Biology and the Environment, Nanjing Forestry University, Nanjing 210037, China
4 Beijing Jingxi Forestry Farm, Beijing 102300, China

Received:2018-12-31 Revised:2019-05-30 Online:2019-06-20 Published:2019-09-30
Contact: XU Jun-Liang
Supported by:
Supported by the National Natural Science Foundation of China(41401063);Supported by the National Natural Science Foundation of China(41801026);the China Scholarship Council(201808410575);the China Scholarship Council(201908410061)

摘要/Abstract

摘要：

枝条是碳供应器官和碳需求器官的连接者, 研究其非结构性碳水化合物(NSC)含量的季节变化对理解树木体内的碳分配至关重要。该研究以秦岭东段栓皮栎(Quercus variabilis)优势群落为研究对象, 于2016年5月至2017年5月, 在其分布的海拔上下限(650 m和970 m), 通过在展叶期采用旬尺度和在非展叶期采用月尺度相结合的周期性取样方法(共计12次), 测定栓皮栎枝条NSC组分及含量, 并观测同期叶片物候变化。结果表明: (1)栓皮栎枝条NSC含量随季节波动较小, 变化差异不显著。但枝条可溶性糖含量(高海拔)或淀粉含量(低海拔)在一定生境条件下, 均存在明显的季节波动, 说明栓皮栎枝条可溶性糖和淀粉之间存在动态转化过程。(2)栓皮栎枝条NSC组成以可溶性糖为主(61%), 这可能是该树种在暖温带季风气候区所采取的生长策略。(3)土壤含水量(正相关)和饱和水汽压差(负相关)分别是在高海拔和低海拔影响栓皮栎枝条NSC含量的主导环境因子, 说明相比高海拔, 低海拔的栓皮栎可能对高温引起的水分胁迫更敏感。(4)结合叶片物候发现, 栓皮栎枝条NSC含量最大值出现在萌芽前(3月中下旬, 11%左右), 最小值出现在展叶后期(4月末, 5%左右), 叶片萌芽展叶后枝条NSC含量下降。总体而言, 枝条NSC含量在高低海拔不存在显著差异, 但春季萌芽前后存在显著差异, 海拔引起的叶片物候时间差极可能是造成这一现象的主要原因。研究结果说明, 栓皮栎叶片物候会直接影响枝条NSC含量的季节变化, 枝条NSC含量对叶片萌芽生长至关重要, 研究结果有助于加深对栓皮栎树体内碳调配机制的理解。

关键词: 非结构性碳水化合物, 栓皮栎, 叶片物候, 海拔, 饱和水汽压差

Abstract:

Aims Measure of non-structural carbohydrate (NSC) reserves indicates tree carbon surplus or shortage stored. Branches connect NSC sources (leaves) and NSC sinks (stemwood, root) of woody plants. Therefore, the seasonal dynamics of NSC concentration in branches will be of important implications for understanding and modeling plant carbon allocation. Methods We conducted a field survey monitoring branch NSC concentrations of Chinese cork oak (Quercus variabilis). We also synchronously observed the leaf phenology of the trees in uneven-aged secondary oak forests at its upper and lower distribution limits (650 m to 970 m) in east Qinling Mountain ranges. Sampling intervals were set semimonthly/monthly during the leaf unfolding period (March to May), and monthly/bimonthly during the tree’s full growing season (June to November) from May 2016 to May 2017. Important findings (1) The NSC measures in the tree branches had weak seasonal changes at both sites. However, the soluble sugar (SS) concentrations at the upper elevation site and the starch (S) concentrations at the lower site had significant seasonal changes. The relative stable NSC levels vs. larger seasonal oscillations of soluble sugar and starch may be explained by the mutual conversion between soluble sugar and starch in the tree branches. (2) Soluble sugar was the major contributor to the total NSC in oak branches, accounting for approximately 61% of it. Here the soluble sugars performed as quick C whereas starch acted mostly as reserved C for future use, it could be inferred that the Q. variabilis, a warm temperate deciduous tree species, developed this feature as its life strategy to survive in warm temperate climate. (3) Soil water availability was positively related to the NSC measures at the high elevation site, while vapor pressure deficit (VPD) was negatively related to the NSC at the low elevation site, indicating the oak may be more drought-susceptible to water stress at lower elevation. (4) The maximum and the minimum concentrations of NSC in tree branches were observed before bud break (late March, approximately 11%) and when full leaf expansion (late April, approximately 11%), respectively. These extremes of NSC could be partly explained by the simultaneous leaf phenological dynamics. Considering the fact of carbon supply for bud break and leaf development via branches rather than by photosynthesis, it was reasonable that the NSC concentration in branches of Q. variabilis reached its maximum before the bud break, and did not change significantly with elevation. Not surprisingly, the significant differences in branches NSC with elevation only occurred during bud break in spring, as a later phenology occurred at higher elevation. In conclusion, this study confirms that the leaf phenology of Q. variabilis directly affects the branches seasonal patterns of NSC, particularly in the spring. The relationship between the carbon budget of the oak branches and its aboveground phenology should be more emphasized for further comprehension on the NSC metabolism.

Key words: non-structural carbohydrate, Quercus variabilis, leaf phenology, altitude, vapor pressure deficit

章异平, 海旭莹, 徐军亮, 吴文霞, 曹鹏鹤, 安文静. 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. 植物生态学报, 2019, 43(6): 521-531. DOI: 10.17521/cjpe.2018.0325

ZHANG Yi-Ping, HAI Xu-Ying, XU Jun-Liang, WU Wen-Xia, CAO Peng-He, AN Wen-Jing. Seasonal dynamics of non-structural carbohydrate content in branch of Quercus variabilis growing in east Qinling Mountain range. Chinese Journal of Plant Ecology, 2019, 43(6): 521-531. DOI: 10.17521/cjpe.2018.0325

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0325

https://www.plant-ecology.com/CN/Y2019/V43/I6/521

图/表 8

表1 秦岭东段栓皮栎林2016和2017年高低海拔环境因子月值

Table 1 Monthly measured environmental parameters at upper and lower altitudes in Quercus variabilis forests in the east Qinling Mountain range in 2016 and 2017

年 Year	月 Month	空气温度 Air temperature (℃)		土壤温度 Soil temperature (℃)		相对湿度 Relative humidity (%)		土壤体积含水量 SWC (%)
年 Year	月 Month	高海拔 Upper	低海拔 Lower	高海拔 Upper	低海拔 Lower	高海拔 Upper	低海拔 Lower	高海拔 Upper	低海拔 Lower
2016	3	8.2	8.9	5.6	7.0	47	51	16.6	15.4
	4	14.7	15.1	10.8	13.2	62	69	17.9	15.9
	5	16.5	16.8	13.5	15.6	65	71	14.8	11.9
	6	22.0	22.2	18.1	20.4	72	77	17.8	14.2
	7	23.5	24.0	20.4	23.2	84	86	16.7	13.1
	8	22.6	23.3	20.9	23.6	87	90	16.5	13.2
	9	18.6	19.5	18.1	20.1	77	80	12.0	11.0
	10	11.4	12.6	13.6	14.9	89	90	13.8	12.5
	11	6.1	6.7	7.6	8.8	74	79	20.6	17.4
	12	2.0	3.3	2.6	6.0	74	76	21.6	NA
2017	1	-0.7	0.1	1.0	1.0	73	70	21.2	12.8
	2	3.1	3.9	NA	NA	67	67	NA	NA
	3	6.3	7.2	5.3	6.4	61	64	20.5	11.9
	4	14.1	14.5	10.3	11.7	59	63	20.7	12.9
	5	18.7	19.1	13.9	15.7	57	63	18.4	10.0

表2 秦岭东段栓皮栎林样地基本情况

Table 2 Physical (altitude, slope and aspect) and forest (canopy closure, average tree age, average tree height and average DBH) measures of sample plots in Quercus variabilis forest in the east Qinling Mountain range

样地 Plot	海拔 Altitude (m)	坡度 Slope (°)	坡向 Aspect	林分郁闭度 Canopy density	平均树龄 Average tree age (a)	平均树高 Average tree height (m)	平均胸径 Average diameter at breast height (cm)
低海拔 Lower altitude	650	25	阳坡 South-facing slope	0.9	26	16	16
高海拔 Upper altitude	970	28	阳坡 South-facing slope	0.8	30	15	16

图1 秦岭东段高海拔栓皮栎枝条非结构性碳水化合物(NSC)及其组分含量的时间动态变化(平均值±标准偏差)。图中阴影代表栓皮栎非生长季(基于叶片物候)。

Fig. 1 Chronological changes of non-structural carbohydrate (NSC) and its components in the branches of Quercus variabilis at upper altitude in east Qinling Mountain (mean ± SD). Shaded area represent the non-growing season of Q. variabilis based on leaf phenology.

图2 秦岭东段低海拔栓皮栎枝条非结构性碳水化合物(NSC)及其组分含量的时间动态变化(平均值±标准偏差)。图中阴影代表栓皮栎非生长季(基于叶片物候)。

Fig. 2 Chronological changes of non-structural carbohydrate (NSC) and its components in the branches of Quercus variabilis at lower altitude in east Qinling Mountain (mean ± SD). Shaded area represent the non-growing season of Q. variabilis based on leaf phenology.

图3 秦岭东段不同海拔栓皮栎枝条非结构性碳水化合物(NSC)及其组分含量对比分析(平均值±标准偏差)。不同小写字母表示同一采样日不同海拔间NSC及其组分差异显著(p < 0.05, Paired-sample t test), 其中NSC用字母ab表示, 淀粉用字母cd表示, 可溶性糖用字母ef表示。

Fig. 3 A comparison of non-structural carbohydrate (NSC) and its components in branches of Quercus variabilis at different altitudes in east Qinling Mountain (mean ± SD). Different lowercase letters indicate significant differences between two altitudes based on Paired-sample t test (p < 0 .05). Letters a and b refer to NSC, letters c and d refer to starch, and letters e and f refer to soluble sugars.

表3 秦岭东段不同海拔环境因子与栓皮栎枝条非结构性碳水化合物(NSC)含量相关性分析

Table 3 Pearson correlation between measured environmental variables and branch NSC contents of Quercus variabilis at upper and lower altitudes in east Qinling Mountain

样地 Plot		空气温度 T_a (℃)	土壤温度 T_s (℃)	相对湿度 RH (%)	土壤体积含水量 SWC (%)	降水量 P (mm)	饱和水汽压差 VPD (kPa)
高海拔 Upper altitude	SS	-0.578	-0.589	-0.164	0.717^*	-0.190	-0.278
	S	-0.190	0.008	0.121	-0.537	0.049	-0.185
	NSC	-0.590	-0.472	-0.055	0.235	-0.123	-0.349
	SS/S	-0.178	-0.194	0.121	0.644^*	-0.261	-0.297
低海拔 Lower altitude	SS	-0.591	-0.453	0.378	0.315	-0.051	-0.552
	S	-0.343	-0.250	0.722^*	0.302	-0.470	-0.788^**
	NSC	-0.564	-0.423	0.703^*	0.382	-0.349	-0.845^**
	SS/S	0.296	0.174	-0.546	-0.246	0.821^**	0.702^**

图4 秦岭东段高低海拔栓皮栎叶片物候实测值及Gompertz方程拟合曲线。▲, 高海拔叶片物候实测值; ■, 低海拔叶片物候实测值; ┅, 高海拔叶片物候拟合线(R2 > 0.99); ━, 低海拔叶片物候拟合线(R2 > 0.99)。

Fig. 4 A comparison between measured and modeling values extracted from Gompertz fitting curve of leaf phenology of uppnr and lower altitude Quercus variabilis in east Qinling Mountain. ▲, measured value of leaf phenology at upper altitude; ■, measured value of leaf phenology at lower altitude; ┅, gompertz fitting curve of leaf phenology of upper altitude; ━, gompertz fitting curve of leaf phenology of lower altitude.

表4 秦岭东段栓皮栎叶片物候主要阶段日期(拟合值)

Table 4 The dates of main leaf phenology phase of Quercus variabilis based on Gompertz fitting values in east Qinling Mountain

样地 Plot	展叶初期 Onset leaf expansion	展叶中期 Mid-stage leaf expansion	展叶末期 Leaf expansion completed
高海拔 Upper altitude	04-12	04-22	05-10
低海拔 Lower altitude	04-09	04-19	05-07

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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

Seasonal dynamics of non-structural carbohydrate content in branch of Quercus variabilis growing in east Qinling Mountain range

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