云南轿子山不同海拔急尖长苞冷杉径向生长动态及其低温阈值

doi:10.17521/cjpe.2021.0399

植物生态学报 ›› 2022, Vol. 46 ›› Issue (9): 1038-1049.DOI: 10.17521/cjpe.2021.0399

云南轿子山不同海拔急尖长苞冷杉径向生长动态及其低温阈值

朱明阳¹, 林琳¹, 佘雨龙¹, 肖城材¹, 赵通兴¹, 胡春相², 赵昌佑², 王文礼¹^,^*()

¹云南大学生态与环境学院, 昆明 650500
²云南轿子山国家级自然保护区管护局, 昆明 651500

收稿日期:2021-11-08 接受日期:2021-12-13 出版日期:2022-09-20 发布日期:2022-10-19
通讯作者: 王文礼
作者简介:*王文礼:ORCID:0000-0003-1536-9890 (wwl@ynu.edu.cn)
基金资助:
国家自然科学基金(32160354);国家自然科学基金(31500174)

Radial growth and its low-temperature threshold of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan, China

ZHU Ming-Yang¹, LIN Lin¹, SHE Yu-Long¹, XIAO Cheng-Cai¹, ZHAO Tong-Xing¹, HU Chun-Xiang², ZHAO Chang-You², WANG Wen-Li¹^,^*()

¹School of Ecology and Environment Science, Yunnan University, Kunming 650500, China
²Jiaozishan National Nature Reserve, Kunming 651500, China

Received:2021-11-08 Accepted:2021-12-13 Online:2022-09-20 Published:2022-10-19
Contact: WANG Wen-Li
Supported by:
National Natural Science Foundation of China(32160354);National Natural Science Foundation of China(31500174)

摘要/Abstract

摘要：

环境对树木的功能和生存具有至关重要的意义, 但环境因素影响树木发育的过程仍有待探究。树木径向生长动态监测是深刻了解树木生长状况及其对气候变化响应的重要手段。该研究以云南轿子山不同海拔树线树种急尖长苞冷杉(Abies georgei var. smithii)为研究对象, 对其年内径向生长动态进行监测, 以期明晰树木径向生长各阶段起止时间的海拔差异, 分析不同海拔树木形成层活动和木质部分化对温度的响应, 辨析树木径向生长的低温阈值。结果表明: (1)海拔越高, 径向生长开始的时间越晚, 结束越早, 生长季缩短。海拔每上升100 m, 急尖长苞冷杉径向生长的开始时间推迟4.7 d, 结束时间提前7.2 d, 生长季缩短12.8 d; (2) 3个海拔急尖长苞冷杉的径向生长具有相似的低温阈值(约5 ℃), 温度控制形成层活动的开始与木质部分化的结束; (3)不同海拔急尖长苞冷杉形成层的细胞分裂活动均在温度较高时(夏至日前后)减弱。形成层活动的结束与海拔引起的温度变化关系较弱, 光周期可能参与了形成层活动结束的调控, 以确保当年新生细胞能够在冻害来临前完成木质化。该研究结果有助于加深树木生长动态对气候变化响应机制的认识, 为更好地适应和应对气候变化提供科学依据。

关键词: 径向生长, 温度阈值, 形成层活动, 木质部分化, 微树芯

Abstract:

Aims In view of the overarching significance for tree function and survival, it is essential to explore the environmental influences on tree development. Our objective was to clarify the altitudinal differences in the onset and offset of each stage of tree radial growth, to analyze the response of cambium activity and xylem differentiation to temperature at different altitudes, and to identify low temperature threshold for radial growth of trees.

Methods Weekly microcores and hourly in situ climate data were collected at 3 600, 3 800 and 4 000 m along the altitude gradient of Abies georgei var. smithii forest in Jiaozi Mountain, Yunnan.

Important findings The results showed that: (1) There were differences in the phenology of radial growth at three altitudes. The onset of radial growth was delayed by 4.7 d per 100 m, the end of radial growth was advanced by 7.2 d per 100 m, and the growing season was shortened by 12.8 d per 100 m with increasing altitude; (2) The radial growth of Abies georgei var. smithii at different altitudes had similar low temperature thresholds (about 5 °C), and temperature controlled the onset of cambium activity and the end of xylem differentiation; (3) The degree of cell division activity at different altitude decreased at higher temperatures (around the summer solstice), and photoperiod was involved in the regulation of the end of cambium activity to ensure the completion of cell mature before the freezing. The relationship between the cambium activity and altitude-induced temperature change is weak. The photoperiod may be involved in regulating the end of cambium activity to ensure the completion of cell mature before the freezing. This study contributes to understand the response mechanisms of tree growth dynamics to climate change and provides a scientific basis for better adaptation and response to climate change.

Key words: radial growth, temperature threshold, cambium activity, xylem differentiation, micro-core

朱明阳, 林琳, 佘雨龙, 肖城材, 赵通兴, 胡春相, 赵昌佑, 王文礼. 云南轿子山不同海拔急尖长苞冷杉径向生长动态及其低温阈值. 植物生态学报, 2022, 46(9): 1038-1049. DOI: 10.17521/cjpe.2021.0399

ZHU Ming-Yang, LIN Lin, SHE Yu-Long, XIAO Cheng-Cai, ZHAO Tong-Xing, HU Chun-Xiang, ZHAO Chang-You, WANG Wen-Li. Radial growth and its low-temperature threshold of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan, China. Chinese Journal of Plant Ecology, 2022, 46(9): 1038-1049. DOI: 10.17521/cjpe.2021.0399

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0399

https://www.plant-ecology.com/CN/Y2022/V46/I9/1038

图/表 15

图1 云南轿子山研究区域示意图。

Fig. 1 Schematic diagram of the study area in Jiaozi Mountain, Yunnan.

表1 云南轿子山不同海拔急尖长苞冷杉样树特征

Table 1 Sample trees characteristics of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan

海拔 Altitude (m)	编号 No.	基径 Base diameter (cm)	胸径 Diameter at breast (cm)	高度 Height (m)	树龄 Tree age (a)
3 600	1	32	25	13.0	58
	2	21	17	11.0	27
	3	23	18	12.0	45
	4	20	16	12.0	51
	5	22	17	12.0	40
3 800	6	45	35	12.0	32
	7	32	25	11.0	24
	8	16	12	12.0	20
	9	34	27	13.0	24
	10	22	17	10.0	18
4 000	11	20	17	5.1	27
	12	57	44	6.8	55
	13	19	15	5.0	27
	14	19	15	3.9	22
	15	21	16	5.2	20

图2 云南轿子山急尖长苞冷杉径向生长各阶段示意图。CZ, 休眠形成层; EN, 细胞增大; M, 细胞成熟; WT, 细胞壁增厚。

Fig. 2 Schematic diagram of each stage of radial growth of Abies georgei var. smithii in Jiaozi Mountain, Yunnan. CZ, dormant cambium; EN, cell enlargement; M, cell maturation; WT, cell wall-thickening.

图3 2020年云南轿子山不同海拔样地微生境平均温度变化趋势图。

Fig. 3 Mean temperature trends of microhabitats at different altitude sample sites in Jiaozi Mountain, Yunnan in 2020.

表2 2020年云南轿子山不同海拔样地内空气、土壤年平均温度、最高温度、最低温和直减率(平均值±标准误)

Table 2 Mean, maximum, minimum temperature and lapse rate (mean ± SE) of air and soil at different altitudes in Jiaozi Mountain, Yunnan in 2020

	空气温度 Air temperature (℃)				土壤温度 Soil temperature (℃)
海拔 Altitude (m)	平均 Mean	最高 Maximum	最低 Minimum	直减率 Direct reduction rate (℃·m^-1)	平均 Mean	最高 Maximum	最低 Minimum	直减率 Direct reduction rate (℃·m^-1)
3 600	5.56	14.25	-4.66	0.005 9 ± 0.000 7	5.82	11.26	-0.39	0.005 7 ± 0.000 6
3 800	4.25	10.06	-7.49		4.57	11.30	-3.73
4 000	3.20	10.41	-9.04		3.54	9.33	-1.10

图4 2020年云南轿子山不同海拔样地空气、土壤温度日较差。

Fig. 4 Daily range of air and soil temperature at different altitude sample sites in Jiaozi Mountain, Yunnan in 2020.

图5 2020年云南轿子山不同海拔急尖长苞冷杉径向生长各发育阶段的起止时间(平均值±标准误)。

Fig. 5 Start and end time of each development stage of radial growth of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan in 2020 (mean ± SE).

图6 2020年云南轿子山不同海拔急尖长苞冷杉径向生长各发育阶段细胞数随时间变化图(平均值±标准误)。

Fig. 6 Variation of cell number in each development stage with time of radial growth of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan in 2020 (mean ± SE).

图7 2020年云南轿子山急尖长苞冷杉径向生长各阶段生长时间所占比例。CZ, 形成层; EN, 细胞膨大; M, 成熟细胞; WT, 细胞壁加厚。

Fig. 7 Proportion of growth time in each stage of radial growth of Abies georgei var. smithii in Jiaozi Mountain, Yunnan in 2020. CZ, cambium; EN, enlargement; M, maturation; WT, wall-thickening.

表3 2020年云南轿子山不同海拔急尖长苞冷杉径向生长各阶段开始、结束以及持续时间单因素方差分析

Table 3 One-way ANOVA for the onset, ending and duration of each stage of radial growth of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan in 2020

	形成层细胞 Cambial cell	细胞膨大 Cell enlarging	细胞壁加厚 Cell wall-thickening	细胞成熟 Mature cells
开始 Onset	0.001^**	0.001^**	0.016^*	0.826
结束 Ending	0.534	0.024^*	0.311	0.011^*
持续 Duration	0.096	0.001^**	0.02^*	0.015^*

表4 云南轿子山2020年不同海拔样地生长季空气、土壤温度

Table 4 Air and soil temperature during the growing season at different altitudes sample sites in Jiaozi Mountain, Yunnan in 2020

					空气温度 Air temperature (℃)					土壤温度 Soil temperature (℃)
海拔 Altitude (m)	最低 Minimum	最高 Maximum	平均 Mean	>0 ℃积温 >0 ℃ accumulation	>5 ℃积温 >5 ℃ accumulation	最低 Minimum	最高 Maximum	平均 Mean	>0 ℃积温 >0 ℃ accumulation	>5 ℃积温 >5 ℃ accumulation	生长季长度 (平均值±标准差) Growing season length (mean ± SD) (d)
3 600	0.06	14.25	8.87	1 775.83	1 730.18	2.80	11.26	8.78	1 718.75	1 654.99	195 ± 4
3 800	1.52	10.06	7.63	1 419.37	1 389.24	2.79	11.30	7.90	1 457.26	1 420.16	177 ± 1
4 000	2.40	10.41	7.19	1 000.10	959.21	2.70	9.33	7.33	983.20	927.57	144 ± 16

表5 2020年云南轿子山不同海拔急尖长苞冷杉的生长量和生长速率(平均值±标准误)

Table 5 Radial growth volume and rate of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan in 2020 (mean ± SE)

海拔 Altitude (m)	细胞总数 Total number of cells	最大生长速率 Maximum growth rate (cell·d^-1)	加速拐点 Accelerated inflection point (d)	平均生长速率 Mean growth rate (cell·d^-1)	n
3 600	75.79 ± 3.86	0.64 ± 0.03	159.90 ± 2.70	0.39 ± 0.02	5
3 800	58.81 ± 3.55	0.67 ± 0.03	157.63 ± 3.26	0.41 ± 0.02	5
4 000	82.68 ± 9.13	0.76 ± 0.11	171.33 ± 9.00	0.47 ± 0.07	5
p	0.008	0.147	0.055	0.147

图8 2020年云南轿子山急尖长苞冷杉不同海拔径向生长的Gompertz生长曲线(平均值±标准误)。

Fig. 8 Gompertz growth curve for radial growth of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan in 2020 (mean ± SE).

图9 2020年云南轿子山急尖长苞冷杉径向生长开始和结束与温度变量的皮尔逊相关指数。ADDS > 0 ℃, 大于0 ℃的空气积温; ADDS > 5 ℃, 大于5 ℃的空气积温; SDDS > 0 ℃, 大于0 ℃的土壤积温; SDDS > 5 ℃, 大于5 ℃的土壤积温; Tamax, 最高气温; Tamean, 平均气温; Tamin, 最低气温; Tsmax, 最高土壤温度; Tsmean, 平均土壤温度; Tsmin, 最低土壤温度。

Fig. 9 Pearson’s correlation index of radial growth onset and end of radial growth of Abies georgei var. smithii with temperature variables in 2020. ADDS > 0 °C, air accumulation temperature greater than 0 °C; ADDS > 5 °C, air accumulation temperature greater than 5 °C; SDDS > 0 °C, soil accumulation temperature greater than 0 °C; SDDS > 5 °C, soil accumulation temperature greater than 5 °C; Tamax, maximum air temperature; Tamean, average air temperature; Tamin, minimum air temperature; Tsmax, maximum soil temperature; Tsmean, average soil temperature; Tsmin, minimum soil temperature.

表6 2020年云南轿子山急尖长苞冷杉径向生长的温度阈值(平均值±标准误)

Table 6 Temperature thresholds for the onset and end of radial growth of Abies georgei var. smithii (mean ± SE) in 2020

	温度变量 Temperature variables	径向生长开始 Radial growth onset	n	p	径向生长结束 End of radial growth	n	p	形成层活动结束 End of cambium active	n	p
空气温度 Air temperature (℃)	平均 Mean	4.78 ± 0.13	12	0.81	5.13 ± 0.20	12	0.14	8.38 ± 0.57	12	0.64
	最高 Maximum	6.91 ± 0.23	12	0.00^**	6.98 ± 0.38	12	0.00^**	16.27 ± 8.01	12	0.38
	最低 Minimum	2.35 ± 0.31	12	0.00^**	3.27 ± 0.35	12	0.00^**	5.65 ± 0.30	12	0.02^*
土壤温度 Soil temperature (℃)	平均 Mean	4.02 ± 0.19	12	0.10	4.96 ± 0.19	12	0.14	8.07 ± 0.50	12	0.68
	最高 Maximum	5.20 ± 0.26	12	0.00^**	6.33 ± 0.21	12	0.01^*	8.44 ± 0.57	12	0.17
	最低 Minimum	2.58 ± 0.22	12	0.00^**	3.70 ± 0.41	12	0.00^**	7.81 ± 0.46	12	0.26

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云南轿子山不同海拔急尖长苞冷杉径向生长动态及其低温阈值

Radial growth and its low-temperature threshold of Abies georgei var. smithii at different altitudes in Jiaozi Mountain, Yunnan, China

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