气候变暖对陆生植物的影响

温度是影响植物生长、发育和功能的重要环境因子,是调节许多陆地生态系统生物地球化学过程的关键因素之一.全球气候持续变暖直接或间接地对陆生植物产生不同程度的影响.本文从不同方面分析了植物对温度升高及其它生态环境因子变化交互作用下的生理生态适应机制,包括生态系统的土壤呼吸、植物的气体交换、水分关系、生物量和生产力的响应等方面的研究进展,并就未来开展陆生植物对气候变暖响应研究提出加强和改进的设想.     

3,4-二羟基苯乙酮胁迫对天山云杉种子萌发过程中内源植物激素含量变化的影响

3,4-二羟基苯乙酮(DHAP)是天山云杉(Picea schrenkiana ssp. tianschanica)叶和凋落物中存在的主要自毒物质, 是导致天山云杉林天然更新障碍的原因之一。为了解释自毒物质发生作用的生理机制, 该文设计多个浓度梯度的DHAP溶液处理天山云杉种子, 以发芽率和发芽势为种子萌发参数, 运用反相超高效液相色谱(UPLC)分析技术, 检测了种子萌发过程中内源植物激素玉米素(ZT)、赤霉素(GA₃)、吲哚乙酸(IAA)和脱落酸(ABA)含量水平的变化。研究结果表明: DHAP处理对天山云杉种子萌发影响具有浓度效应, 表现为5.0 mmol·L^-1 > 0.1 mmol·L^-1 > 1.0 mmol·L^-1 >对照, 即5.0 mmol·L^-1 DHAP处理组对种子萌发的抑制作用最强、0.1 mmol·L^-1 DHAP处理组次之、1.0 mmol·L^-1 DHAP处理组最弱; DHAP处理组的种子内源ZT、GA₃浓度水平降低, ABA含量升高, GA₃浓度峰值出现时间延迟, IAA浓度在高浓度(5.0 mmol·L^-1 DHAP)处理组短时间内(3-6天)过量积累, 1.0和5.0 mmol·L^-1 DHAP处理组的种子内源ABA浓度峰值出现时间延迟; DHAP处理种子萌发1-6天时, ZT/(GA₃+IAA)比值降低, IAA/ZT、ABA/ZT比值增大; ABA/(ZT + GA₃ + IAA)比值在0.1 mmol·L^-1 DHAP处理组增大, 在5.0 mmol·L^-1 DHAP处理组降低。DHAP处理引发种子内源激素含量水平及激素含量间比值变化, 可能是抑制、延迟天山云杉种子萌发的主要原因。     

苔藓植物对植物天然更新的影响

种子是天然更新的基础,幼苗和幼树的生长是天然更新过程中最敏感的阶段。苔藓植物作为许多生态系统的主要地被物,在维管植物天然更新过程中发挥着不容忽视的作用。在许多植被类型中,都有发育良好的苔藓群落,大量的研究证明它们的存在影响到维管植物的萌发、建成和生长,并且可能进一步影响到物种的共生,有一些种类甚至可能指示树种的更新。本文就苔藓植物对物种更新早期阶段的作用和影响进行了综述。包括苔藓植物的生理特性造成的微气候变化;苔藓群落对种子传播、萌发及幼苗建成的影响。并讨论了苔藓植物与更新物种的种间关系及其对幼苗生长的化感作用。     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

Responses of white spruce (Picea glauca) to experimental warming at a subarctic alpine treeline

From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open‐top chambers (OTCs) distributed equally between opposing north and south‐facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1.8°C and annual growing degree‐days by one‐third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1.0°C warmer and the snow‐free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem‐II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure‐related differences. Our results suggest that the growth of seedlings on north‐facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south‐facing slopes appears limited by winter desiccation and cold‐induced photoinhibition.     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014-2016年, 该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗: >150 m ², 中林窗: 100-150 m ², 小林窗: 50-100 m ², 对照: 林下)下进行栲(Castanopsis fargesii)种子野外播种实验, 并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明: (1)栲幼苗出土时间从7月持续到12月, 出苗时间较长, 大林窗对幼苗出土具有延迟作用; (2)栲种子野外平均萌发率为(62.8 ± 2.0)%, 第3个生长季(2016年)末幼苗平均存活率为(65.1 ± 2.2)%, 枯萎是栲幼苗死亡的主要原因; (3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响, 对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响; (4)林冠环境在第1个生长季对幼苗生长无明显影响, 但在第2个和第3个生长季具有显著影响, 大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下, 比叶面积显著低于林下; (5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高, 根质量比和根冠比降低, 并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下, 根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量, 受林冠条件影响较弱, 后期则依赖光合作用, 受林冠条件影响较强, 从整个幼苗建成过程看, 大、中林窗更有利于栲幼苗定居。     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

升温对祁连山东部青海云杉径向生长的影响

在祁连山东部西营河流域不同海拔采集年轮样品,利用树木年代学方法建立高海拔(3000 m)、中海拔(2750 m)、低海拔(2500 m)青海云杉标准化年表,将年轮宽度指数与不同时段气温和降水分别进行相关分析,研究祁连山东部不同海拔青海云杉径向生长对气候变暖的响应。结果表明: 水热是祁连山东部青海云杉径向生长的主要限制因子,不同海拔树木的生长限制因子基本一致。在显著升温之前(1961—1986年),低、中、高海拔3个样点的青海云杉径向生长均与上一年7—8月和当年8月平均最高温呈显著负相关,与当年6月相对湿度呈显著正相关。显著升温之后(1986—2014年),高、中、低海拔树木生长与气温仍呈显著负相关,而与当年2月的相对湿度由升温前的不显著负相关转为显著正相关,与6月降水和相对湿度由升温前的显著正相关转为不显著的负相关。升温导致祁连山东部各海拔青海云杉树轮生长变慢,其中高海拔所受影响最大。气候变暖导致的干旱胁迫可能是青海云杉径向生长发生变化的主要原因。     

Altered dynamics of forest recovery under a changing climate

Forest regeneration following disturbance is a key ecological process, influencing forest structure and function, species assemblages, and ecosystem–climate interactions. Climate change may alter forest recovery dynamics or even prevent recovery, triggering feedbacks to the climate system, altering regional biodiversity, and affecting the ecosystem services provided by forests. Multiple lines of evidence – including global‐scale patterns in forest recovery dynamics; forest responses to experimental manipulation of CO₂, temperature, and precipitation; forest responses to the climate change that has already occurred; ecological theory; and ecosystem and earth system models – all indicate that the dynamics of forest recovery are sensitive to climate. However, synthetic understanding of how atmospheric CO₂ and climate shape trajectories of forest recovery is lacking. Here, we review these separate lines of evidence, which together demonstrate that the dynamics of forest recovery are being impacted by increasing atmospheric CO₂ and changing climate. Rates of forest recovery generally increase with CO₂, temperature, and water availability. Drought reduces growth and live biomass in forests of all ages, having a particularly strong effect on seedling recruitment and survival. Responses of individual trees and whole‐forest ecosystems to CO₂ and climate manipulations often vary by age, implying that forests of different ages will respond differently to climate change. Furthermore, species within a community typically exhibit differential responses to CO₂ and climate, and altered community dynamics can have important consequences for ecosystem function. Age‐ and species‐dependent responses provide a mechanism by which climate change may push some forests past critical thresholds such that they fail to recover to their previous state following disturbance. Altered dynamics of forest recovery will result in positive and negative feedbacks to climate change. Future research on this topic and corresponding improvements to earth system models will be a key to understanding the future of forests and their feedbacks to the climate system.     

Effects of warming on seed germination of woody species in temperate secondary forests

• Seed germination, a critical stage of the plant life cycle providing a link between seeds and seedlings, is commonly temperature-dependent. The global average surface temperature is expected to rise, but little is known about the responses of seed germination of woody plants in temperate forests to warming.
• In the present study, dried seeds of 23 common woody species in temperate secondary forests were incubated at three temperature sequences without cold stratification and after experiencing cold stratification. We calculated five seed germination indices and the comprehensive membership function value that summarized the above indicators.
• Compared to the control, +2 and +4 °C treatments without cold stratification shortened germination time by 14% and 16% and increased the germination index by 17% and 26%, respectively. For stratified seeds, +4 °C treatment increased germination percentage by 49%, and +4 and +2 °C treatments increased duration of germination and the germination index, and shortened mean germination time by 69%, 458%, 29% and 68%, 110%, 12%, respectively. The germination of Fraxinus rhynchophylla and Larix kaempferi were most sensitive to warming without and with cold stratification, respectively. Seed germination of shrubs was the least sensitive to warming among functional types.
• These findings indicate warming (especially extreme warming) will enhance the seedling recruitment of temperate woody species, primarily via shortening the germination time, particularly for seeds that have undergone cold stratification. In addition, shrubs might narrow their distribution range

     

芦芽山阳坡不同海拔白杄径向生长对气候变暖的响应

使用树轮生态学方法研究了山西芦芽山建群种白杄(Picea meyeri)径向生长对气候变暖的响应状况, 发现随着气温升高, 不同海拔白杄生长与气候因子关系的变化存在差别。研究区气温可以分为1958-1983年的气温降低阶段和1984-2007年的气温升高阶段。由气温降低阶段进入气温升高阶段, 低海拔白杄树轮年表的序列间相关系数和第一主成分解释量均增大, 而高海拔白杄树轮年表的序列间相关系数和第一主成分解释量均减小, 表明气候条件对低海拔白杄生长的影响增强而对高海拔白杄生长的影响减弱。随着气温升高, 不同海拔白杄径向生长与气候因子的关系均出现了变化。1958-1983年, 低海拔(2 060 m)白杄生长与7月降水量显著正相关(p < 0.05), 而在1984-2007年, 这一关系表现为极显著正相关(p < 0.01), 同时与生长季中5-7月平均气温呈现显著负相关(p < 0.05)。海拔2 330 m, 白杄在1958-1983年与7月降水量极显著正相关(p < 0.01), 进入1984-2007年后与气候因子没有显著相关关系。海拔2 440 m, 白杄生长由1958-1983年的与气候因子没有显著相关关系转变为1984-2007年的与上一年10月平均气温显著负相关(p < 0.05)。高海拔(2 540 m)白杄生长在1958-1983年与上一年11月平均气温极显著负相关(p < 0.01), 在1984-2007年与上一年10月、当年1月平均气温和6月降水量均显著负相关(p < 0.05)。滑动相关分析结果表明, 随着气温升高, 低海拔主要气候因子对生长的影响增强, 而高海拔主要气候因子对生长的影响减弱, 这可能成为高海拔白杄生长对气温升高敏感性降低的原因。在气候变暖的驱动下, 海拔引起的白杄生长与气候因子关系的差异发生了变化。     

Shifts in frog size and phenology: Testing predictions of climate change on a widespread anuran using data from prior to rapid climate warming

Changes in body size and breeding phenology have been identified as two major ecological consequences of climate change, yet it remains unclear whether climate acts directly or indirectly on these variables. To better understand the relationship between climate and ecological changes, it is necessary to determine environmental predictors of both size and phenology using data from prior to the onset of rapid climate warming, and then to examine spatially explicit changes in climate, size, and phenology, not just general spatial and temporal trends. We used 100 years of natural history collection data for the wood frog, Lithobates sylvaticus with a range >9 million km², and spatially explicit environmental data to determine the best predictors of size and phenology prior to rapid climate warming (1901–1960). We then tested how closely size and phenology changes predicted by those environmental variables reflected actual changes from 1961 to 2000. Size, phenology, and climate all changed as expected (smaller, earlier, and warmer, respectively) at broad spatial scales across the entire study range. However, while spatially explicit changes in climate variables accurately predicted changes in phenology, they did not accurately predict size changes during recent climate change (1961–2000), contrary to expectations from numerous recent studies. Our results suggest that changes in climate are directly linked to observed phenological shifts. However, the mechanisms driving observed body size changes are yet to be determined, given the less straightforward relationship between size and climate factors examined in this study. We recommend that caution be used in “space‐for‐time” studies where measures of a species’ traits at lower latitudes or elevations are considered representative of those under future projected climate conditions. Future studies should aim to determine mechanisms driving trends in phenology and body size, as well as the impact of climate on population density, which may influence body size.     

Gap phase regeneration of tree species of differing successional status in a humid tropical forest of Kerala,India

Germination, establishment and growth of seedlings of tree speciesPalaquium ellipticum (primary),Actinodaphne malabarica (late secondary) andMacaranga peltata (early secondary) were studied in a humid tropical forest at Nelliampathy, in the Western Ghats of Kerala. While the primary species completed its germination within a brief period of 1.5 months, at the other extreme, early secondary species showed slow germination extending for about 5 months, the late secondary species falling in between. Although, all the species studied showed higher establishment and growth under gaps, the early secondary species were more responsive compared to the primary species. Primary species showed better establishment in undisturbed sites and natural gaps than under selection felled gaps; the reverse was true for late and early secondary species. Survival of seedlings increased with gap size, but sharply declined with gap age. Shoot/root ratio was consistently higher in the early secondary species than in the primary species.     

Critical periods for impact of climate warming on early seedling establishment in subarctic tundra

Climate warming is expected to shift bioclimatic zones and plant species distribution. Yet, few studies have explored whether seedling establishment is a possible bottleneck for future migration and population resilience. We test how warming affects the early stages of seedling establishment in 10 plant species in subarctic tundra. To zoom into the life phases where the effects of warming actually take place, we used a novel approach of breaking down the whole‐season warming effect into full factorial combination of early‐, mid‐, and late‐season warming periods. Seeds were sown in containers placed under field conditions in subarctic heath and were exposed to 3 °C elevation of surface temperature and 30% addition of summer precipitation relative to ambient. Heating was achieved with Free Air Temperature Increase systems. Whole‐season heating reduced germination and establishment, significantly in four out of 10 species. The whole‐season warming effect originated from additive effects of individual periods, although some of the periods had disproportionally stronger influence. Early‐germinating species were susceptible to warming; the critical phases were early summer for germination and mid summer for seedling survival. Graminoids, which emerged later, were less susceptible although some negative effects during late summer were observed. Some species with intermediate germination time were affected by all periods of warming. Addition of water generally could not mitigate the negative effects of whole‐season heating, but at individual species level both strengthening and amelioration of these negative effects were observed. We conclude that summer warming is likely to constrain seedling recruitment in open micro sites, which is a common seed regeneration niche in tundra ecosystem. Importantly, we described both significant temporal and species‐specific variation in the sensitivity of seedling establishment to warming which needs to be taken into consideration when modelling population dynamics and vegetation transitions in a warmer climate.     

Sex‐biased oviposition by a nursery pollinator on a gynodioecious host plant: Implications for breeding system evolution and evolution of mutualism

Dioecy, a breeding system where individual plants are exclusively male or female, has evolved repeatedly. Extensive theory describes when dioecy should arise from hermaphroditism, frequently through gynodioecy, where females and hermaphrodites coexist, and when gynodioecy should be stable. Both pollinators and herbivores often prefer the pollen‐bearing sex, with sex‐specific fitness effects that can affect breeding system evolution. Nursery pollination, where adult insects pollinate flowers but their larvae feed on plant reproductive tissues, is a model for understanding mutualism evolution but could also yield insights into plant breeding system evolution. We studied a recently established nursery pollination interaction between native Hadena ectypa moths and introduced gynodioecious Silene vulgaris plants in North America to assess whether oviposition was biased toward females or hermaphrodites, which traits were associated with oviposition, and the effect of oviposition on host plant fitness. Oviposition was hermaphrodite‐biased and associated with deeper flowers and more stems. Sexual dimorphism in flower depth, a trait also associated with oviposition on the native host plant (Silene stellata), explained the hermaphrodite bias. Egg‐receiving plants experienced more fruit predation than plants that received no eggs, but relatively few fruits were lost, and egg receipt did not significantly alter total fruit production at the plant level. Oviposition did not enhance pollination; egg‐receiving flowers usually failed to expand and produce seeds. Together, our results suggest that H. ectypa oviposition does not exert a large fitness cost on host plants, sex‐biased interactions can emerge from preferences developed on a hermaphroditic host species, and new nursery pollination interactions can arise as negative or neutral rather than as mutualistic for the plant.     

Effects of experimental warming on fungal disease progress in oilseed rape

Global warming will influence the growth and development of both crops and pathogens. The aims of this study were to investigate potential effects of future warming on oilseed rape growth and the epidemiology of the three economically important pathogens Verticillium longisporum, Sclerotinia sclerotiorum, and Leptosphaeria maculans (anamorph: Phoma lingam). We utilized climate chambers and a soil warming facility, where treatments represented regional warming scenarios for Lower Saxony, Germany, by 2050 and 2100, and compared results of both approaches on a thermal time scale by calculating degree‐days (dd) from day of sowing, December 1st and March 1st until sampling, the latter correlating best with disease progress. Regression analysis showed that plant growth and growth stages in spring responded almost linearly to increasing thermal time until 1000–1500 dd. Colonization of plant tissue by V. longisporum showed an exponential increase when exceeding 1300–1500 dd and reaching plant growth stage BBCH 74/75 (pod development). V. longisporum colonization of plants may be advanced, potentially leading to higher inoculum densities after harvest and increased economic importance of this pathogen under future warming. Sclerotia germination of S. sclerotiorum reached its maximum at 600–900 dd. Advance of these critical degree‐days may lead to earlier apothecia production, potentially advancing the infection window, whereas the future importance of S. sclerotiorum may remain constant. Severity of phoma crown canker increased linearly with increasing thermal time, but showed also large variation in response to the warming scenarios, suggesting that factors such as canopy microclimate in fall or leaf shedding over winter may play a bigger role for L. maculans infection and disease severity than higher soil temperatures. Thermal time was a suitable tool to combine and integrate data on biological responses to soil and air temperature increases from climate chamber and field experiments.