高山林线变化的更新受限机制研究进展
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摘要
全球林线位置对气候变暖的响应表现为上升、无变化或下降等截然不同趋势,表明影响林线位置及动态的因子十分复杂,除了较普遍认为的低温调控机制外,还存在其它控制林线位置变化的机制。林线向上迁移开始于种子向林线以上的传播及幼苗在林线以上的定居,这些过程中的限制因子均会影响林线的位移,因此研究更新过程及其限制因子对理解高山林线对气候变化的响应具有重要的科学意义。主要从种子和幼苗两个关键阶段综述高山林线森林更新的研究进展。在种子阶段,夏季积温不足导致种子产量和活力下降,风速过低和浓密灌丛限制种子向林线以上传播,近地表的霜冻/水分胁迫和灌木释放的化感物质会阻碍种子在林线以上萌发。在幼苗阶段,除冬季低温外,生长季内较大的温度日振幅和偶然出现的冻害事件也是导致幼苗死亡的重要原因,而低温环境下的强烈光照引起的低温光抑制会显著降低生长季的光合作用;土壤低温、由土壤温度昼夜变化引起的冻举事件、夏季土壤干旱可能会导致幼苗光合作用下降和死亡率上升;积雪太浅会导致生长季早期幼苗水分供应的严重缺乏,但积雪太深会导致幼苗感染真菌的可能性增加;浓密的灌木和草本植物以及植食动物的啃食也会降低林线以上的幼苗存活率。气候变暖对林线幼苗定居的影响复杂且具有很大不确定性,需要进一步研究气候变暖导致的环境因子变化对林线更新各关键阶段的影响。未来气候变暖无疑会导致生长季起始日提前,结束日推迟,这很可能会增加生长季期间尤其是早期的低温冻害事件,对高山林线树种幼苗的存活具有重要影响。在未来研究中,需要找出定义生长季冻害事件的温度阈值,利用长期气象观测数据分析增温背景下生长季早期冻害事件特征的变化趋势,并进一步开展野外模拟增温实验以深刻理解林线树种的种子萌发和幼苗定居与生长季冻害事件的关系,加强对不同地区林线树种的繁殖策略研究,这将有助于人们进一步理解不同区域林线的形成机制并预测未来气候变化条件下林线的动态变化趋势。
Advances of alpine timberline forests during last century are not ubiquitous worldwide,suggesting additional factors and mechanisms likely affect the response of alpine timberline forests to climate warming. Upward shifts of treelines begin with seed dispersal and germination,and seedling establishment above the treeline and any limiting factors during these processes may affect treeline migration. Therefore,investigation of mechanisms controlling seedling recruitment at alpine treeline will be helpful to elucidate treeline formation and its response to future climate change. We reviewed recent advances in tree seedling recruitment at alpine treelines from the key seed and seedling stages. For the seed stage,the seed quantity and quality generally decreased with the sum temperature during summer; the seed dispersal to elevations above treeline was impeded by low wind speed,dense dwarf shrub and grass cover; the ability of seed germination above thetreeline was impaired by frost and water stresses near the ground. Also,the allelochemical properties of shrubs had negative effects on seed germination. For the seedling stage,large temperature amplitudes and freezing events during the growing season,as well as the extremely low temperature during winter,were important factors affect seedling mortality. Also the low-temperature photoinhibition resulted from the combination of low temperature and high sunlight significantly decreased seedling photosynthesis during the growing season. Besides,frost-heave activity induced by large soil temperature amplitude and soil water deficits during summer impeded seedling establishment at and above the treeline. Snowpack could keep the seedlings away from the extremely low air temperature during the winter and supply snowmelt water in the early growing season. However,too long duration of the snowpack might increase the possibility of fungal infection that promote seedling mortality. Dense shrub and grass cover above the treeline and the presence of herbivores might decrease seedling survival. In all,the influence of climate warming on seedling establishment across the timberline ecotone is complex and uncertain.Further research is needed to explore the exact effects of warmth-induced environmental changes to seedling recruitment at the alpine treeline. Since the beginning of the growing season might advance under scenarios of climate warming,which in turn led to more early-season freezing events at and above the treeline,it is important to define the temperature threshold of freezing events to analyze the relationship between growing-season freezing events and increasing temperature in the future.Based on this threshold,we can further disclose the effects of growing-season freezing events on seedling establishment at alpine treeline,which will be helpful to elucidate treeline formation and predict treeline dynamics under future climate change.
Advances of alpine timberline forests during last century are not ubiquitous worldwide,suggesting additional factors and mechanisms likely affect the response of alpine timberline forests to climate warming. Upward shifts of treelines begin with seed dispersal and germination,and seedling establishment above the treeline and any limiting factors during these processes may affect treeline migration. Therefore,investigation of mechanisms controlling seedling recruitment at alpine treeline will be helpful to elucidate treeline formation and its response to future climate change. We reviewed recent advances in tree seedling recruitment at alpine treelines from the key seed and seedling stages. For the seed stage,the seed quantity and quality generally decreased with the sum temperature during summer; the seed dispersal to elevations above treeline was impeded by low wind speed,dense dwarf shrub and grass cover; the ability of seed germination above thetreeline was impaired by frost and water stresses near the ground. Also,the allelochemical properties of shrubs had negative effects on seed germination. For the seedling stage,large temperature amplitudes and freezing events during the growing season,as well as the extremely low temperature during winter,were important factors affect seedling mortality. Also the low-temperature photoinhibition resulted from the combination of low temperature and high sunlight significantly decreased seedling photosynthesis during the growing season. Besides,frost-heave activity induced by large soil temperature amplitude and soil water deficits during summer impeded seedling establishment at and above the treeline. Snowpack could keep the seedlings away from the extremely low air temperature during the winter and supply snowmelt water in the early growing season. However,too long duration of the snowpack might increase the possibility of fungal infection that promote seedling mortality. Dense shrub and grass cover above the treeline and the presence of herbivores might decrease seedling survival. In all,the influence of climate warming on seedling establishment across the timberline ecotone is complex and uncertain.Further research is needed to explore the exact effects of warmth-induced environmental changes to seedling recruitment at the alpine treeline. Since the beginning of the growing season might advance under scenarios of climate warming,which in turn led to more early-season freezing events at and above the treeline,it is important to define the temperature threshold of freezing events to analyze the relationship between growing-season freezing events and increasing temperature in the future.Based on this threshold,we can further disclose the effects of growing-season freezing events on seedling establishment at alpine treeline,which will be helpful to elucidate treeline formation and predict treeline dynamics under future climate change.
引文
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