冻融过程对荒漠区不同微生境下齿肋赤藓渗透调节物含量和抗氧化酶活力的影响

doi:10.17521/cjpe.2015.0050

摘要/Abstract

摘要：

齿肋赤藓(Syntrichia caninervis)作为典型的耐旱藓类, 广泛分布于世界干旱和半干旱荒漠地区, 是古尔班通古特沙漠生物土壤结皮中的优势藓类植物。该沙漠冬季具有稳定的降雪, 初春的积雪融化为植物的生长提供了良好的水热条件。荒漠藓类植物叶片仅具一层细胞, 对外界环境的变化十分敏感, 有关荒漠藓类植物在冬季和早春地表冻融交替过程中如何适应环境剧烈变化的研究鲜见报道。该研究探讨了生长于3种不同微生境(活灌丛、枯死灌丛和裸露地)下的齿肋赤藓, 经由冬季低温冻结到早春融雪复水再到春季中旬自然干燥过程中的生理生化变化特征。结果表明: 不同冻融期、微生境及二者的交互作用能够显著影响齿肋赤藓的渗透调节物质(游离脯氨酸、可溶性糖、可溶性蛋白)含量、丙二醛(MDA)含量、抗氧化酶活性(过氧化氢酶(CAT)、过氧化物酶(POD)和超氧化物歧化酶(SOD)活性)。冬季低温冻结期的极端低温和春季中旬的干燥环境使得齿肋赤藓可溶性糖和MDA含量, 以及3种抗氧化酶活性均显著高于早春融雪期, 可溶性蛋白含量显著低于融雪期。同时, 在融雪期灌丛遮阴所形成的“冷岛效应”使得生长于灌丛下的齿肋赤藓植株可溶性糖和MDA含量显著高于枯死灌丛和裸露地。但灌丛的存在也为春季中旬干旱无雨期齿肋赤藓提供了一个相对良好(含水量高)的生活环境, 其下齿肋赤藓的渗透调节物含量和抗氧化酶活性均显著低于其他两个生境。在整个冻融过程中裸露地齿肋赤藓的3种抗氧化酶活性均显著高于灌丛下, 这可能是由于生活于裸露地的苔藓有更强的耐胁迫特性。

关键词: 苔藓, 生物土壤结皮, 抗氧化酶, 渗透调节物质, 冻融期

Abstract: Aims

Syntrichia caninervis, a typical drought-tolerant moss species found worldwide in various semiarid and arid regions, is the dominant species of soil crust mosses in the Gurbantünggüt Desert, which is a temperate northern desert of Central Asia. It appears able to endure frequent and intensive freeze-thaw cycles in the surface soil and maintain physiological functions active even under extreme low temperatures in winter. However, there have been few studies exploring the adaptive strategies of the species during freeze-thaw processes and interpreting well the phenomena. One of the major goals of this study was to investigate physiological regulatory mechanisms of the species when undergoing freezing-thawing periods and relevant antioxidant enzyme activities.

Methods

A series of microclimate and soil conditions were continuously monitored in the sampling sites starting from November 2013. The samples of the moss were collected during the early spring of 2014 from February to April and from three microhabitats including the sites under live shrub, under dead shrub and on an open ground. The moss samples were quickly treated in the field for keeping freshness and then brought to laboratory for measurement and analysis. The traits/characteristics that reflect physiological and biochemical activities were measured for analyzing responses of the species to freeze-thaw processes and for comparing the impacts of different habitats, which included water content, concentrations of proline, soluble sugar, soluble protein and malondialdehyde (MDA), as well as catalase (CAT), peroxidase (POD), superoxide dismutase (SOD) activities.

Important findings

The results showed that, during different freeze-thaw periods, microhabitats and their interactions with alternative freeze-thaw processes significantly affect the contents of proline, soluble sugar, soluble protein and MDA in shoots of the species, as well as the enzyme activities of CAT, POD, and SOD. Extreme low temperature and drought caused significantly higher soluble sugar and MDA contents, higher CAT, POD and SOD activities, but lower soluble protein content, in snow covered period (February) and dry period (April) than the snow melting period (March). The existence of live shrub decreased temperature around the bushes during snow melting because of the effect of shade, causing increased contents of soluble sugar and MDA in moss shoots, when compared to those growing in the habitats under dead shrub and on open ground. However, live shrub could provide a moister environment for S. caninervis than dead shrub and exposed ground for the duration of desiccation. As the result, the proline content, soluble sugar content and MDA content of the species under the live shrub canopy were the lowest among the three habitats. Furthermore, the mosses lived on the exposed ground showed the highest antioxidant enzyme activities comparing to those under the dead shrub and living shrub. This may suggest that S. caninervis grown on exposed grounds has developed rather a specific adaptations for a harsher environment condition than that occurred in other two habitats.

Key words: bryophytes, biological soil crusts, antioxidant enzymes, osmoregulation substance, freezing-thawing periods

尹本丰, 张元明. 冻融过程对荒漠区不同微生境下齿肋赤藓渗透调节物含量和抗氧化酶活力的影响. 植物生态学报, 2015, 39(5): 517-529. DOI: 10.17521/cjpe.2015.0050

YIN Ben-Feng,ZHANG Yuan-Ming. Impacts of freeze-thaw processes on antioxidant activities and osmolyte contents of Syntrichia caninervis under different desert microhabitats. Chinese Journal of Plant Ecology, 2015, 39(5): 517-529. DOI: 10.17521/cjpe.2015.0050

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0050

https://www.plant-ecology.com/CN/Y2015/V39/I5/517

图/表 9

图1 古尔班通古特沙漠3种不同微生境下生长的齿肋赤藓。A, 裸露地。B, 死灌丛。C, 活灌丛。

Fig. 1 Syntrichia caninervis grown under three microhabitats in Gurbantünggüt Desert. A, Exposed ground. B, Dead shrub. C, Live shrub.

图2 古尔班通古特沙漠三种不同微生境下齿肋赤藓植株。A, 裸露地。B, 死灌丛。C, 活灌丛。

Fig. 2 Syntrichia caninervis shoots collected from three microhabitats in Gurbantünggüt Desert. A, Exposed ground. B, Dead shrub. C, Live shrub.

表1 不同微生境下土壤理化性质的比较(平均值±标准偏差)

Table 1 Comparison of soil physical and chemical properties among different microhabitats (mean ± SD)

项目 Item	活灌丛 Alive shrub	死灌丛 Dead shrub	裸露地 Exposed area
有机碳 Organic matter (g·kg^-1)	3.721 ± 0.318^a	3.898 ± 0.737^a	2.300 ± 0.574^b
全氮 Total nitrogen (g·kg^-1)	0.475 ± 0.073^a	0.495 ± 0.048^a	0.301 ± 0.047^b
全磷 Total phosphorus (g·kg^-1)	0.400 ± 0.014^ab	0.445 ± 0.013^a	0.394 ± 0.059^b
全钾 Total potassium (g·kg^-1)	13.119 ± 0.249^a	12.977 ± 0.127^a	12.194 ± 0.136^b
总盐 Total salt (g·kg^-1)	0.576 ± 0.018^a	0.567 ± 0.079^a	0.514 ± 0.045^a
pH值 pH value	7.936 ± 0.031^a	7.672 ± 0.036^b	7.652 ± 0.262^b
电导率 Electrical conductivity (μs·cm^-1)	0.131 ± 0.009^a	0.129 ± 0.008^a	0.125 ± 0.016^a
冻结期温度(日均温) Temperature in freezing periods (℃)	-9.36	-11.09	-11.67
融雪期温度(日均温) Temperature in snow melting periods (℃)	-2.18	2.81	2.82
春季中旬干旱无雨期温度(日均温) Temperature in dry environment (℃)	9.76	11.64	11.64

图3 冻融期不同微生境下齿肋赤藓植株含水量的比较(平均值±标准偏差)。不同小写字母和大写字母分别表示不同微生境下和不同冻融期齿肋赤藓植株含水量差异显著(p < 0.05)。

Fig. 3 Comparison of water content in Syntrichia caninervis shoots from different microhabitats and freezing-thawing periods (mean ± SD). Different lowercase and capital letters denote statistically significant differences in water content (p < 0.05) between microhabitats and periods, respectively.

表2 不同冻融期不同微生境下齿肋赤藓生化特征重复测量结果的方差分析

Table 2 Repeated measures ANOVA on the effects of microhabitats and Freezing-thawing periods on physiological and biochemical characteristics in Syntrichia caninervis

因子 Factor	脯氨酸含量 Proline content	可溶性糖含量 Soluble sugar content	可溶性蛋白含量 Soluble proteins content	丙二醛含量 MDA content	超氧化物歧化酶活性 SOD activity	过氧化物酶活性 POD activity	过氧化氢酶活性 CAT activity
微生境 Microhabitats	37.695^**	1.841	41.663^**	5.488^*	122.556_**	45.145^**	22.239^**
冻融期 Freezing-thawing periods	266.579^**	919.016^**	325.679^**	1153.946^**	494.308^**	152.878^**	105.681^**
微生境×冻融期 Microhabitats × freezing-thawing periods	40.255^**	27.920^**	3.475^*	7.710^**	50.115^**	15.406^**	3.869^*

图4 冻融期不同微生境下齿肋赤藓脯氨酸、可溶性糖和可溶性蛋白含量的比较(平均值±标准偏差)。不同小写字母和大写字母分别表示不同微生境下和不同冻融期齿肋赤藓渗透调节物质含量差异显著(p < 0.05)。

Fig. 4 Comparison of the contents of compatible solutes (osmoprotectants) in Syntrichia caninervis shoots from different microhabitats and freezing-thawing periods (mean ± SD). Different lowercase and capital letters denote statistically significant differences in solutes (p < 0.05) between microhabitats and periods, respectively.

图5 冻融期不同微生境下齿肋赤藓丙二醛(MDA)含量的比较(平均值±标准偏差)。不同小写字母和大写字母分别表示不同微生境下和不同冻融期齿肋赤藓MDA含量差异显著(p < 0.05)。

Fig. 5 Comparison of the content of malonyldialdehyde (MDA) in Syntrichia caninervis shoots from different microhabitats and freezing-thawing periods (mean ± SD). Different lowercase and capital letters denote statistically significant differences in MDA (p < 0.05) between microhabitats and periods, respectively.

图6 冻融期不同微生境下齿肋赤藓过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性的比较(平均值±标准偏差)。不同小写字母和大写字母分别表示不同微生境下和不同冻融期齿肋赤藓抗氧化酶活性差异显著(p < 0.05)。

Fig. 6 Comparisons of the activities of peroxidase (POD), superoxide dismutase (SOD), catalase (CAT) in Syntrichia caninervis shoots from different microhabitats and freezing-thawing periods (mean ± SD). Different lowercase and capital letters denote statistically significant differences in enzyme activities (p < 0.05) between microhabitats and periods, respectively.

表3 不同微生境下齿肋赤藓生化特征在不同冻融期与温度和含水量之间的相关系数

Table 3 Correlation coefficients among physiological traits in Syntrichhia caninervis shoots and environmental factors in different microhabitats during the freezing-thawing periods

变量 Variable	脯氨酸含量 Proline content	可溶性糖含量 Soluble sugar content	可溶性蛋白含量 Soluble proteins content	丙二醛含量 MDA content	超氧化物歧化酶活性 SOD activity	过氧化物酶活性 POD activity	过氧化氢酶活性 CAT activity
冬季低温冻结期 Freezing period
含水量 Water content	-0.191	0.737^**	0.861^**	-0.051	-0.915^**	-0.800^**	-0.803^**
温度 Temperature	-0.191	0.730^**	0.861^**	0.052	-0.899^**	-0.767^**	-0.797^**
融雪期 Snow melting period
含水量 Water content	0.172	0.462	0.515^*	0.747^**	-0.457	-0.668^**	-0.479
温度 Temperature	-0.008	-0.907^**	-0.696^**	-0.635^*	0.437	0.755^**	0.644^*
春季中旬干旱无雨期 Dry environment
含水量 Water content	-0.812**	-0.676^**	0.747^**	-0.764^**	-0.696^**	-0.758^**	-0.830^**
温度 Temperature	0.802**	0.601^*	-0.688^**	0.707^**	0.710^**	0.767^**	0.809^**

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