植物物候学研究进展

植物物候变化在研究陆地生态系统对气候变化的响应时被誉为"矿井中的金丝雀",全球气候变化愈演愈烈,重新引起了人们对植物物候研究的广泛关注。随着观测技术的发展,在各种空间和生态尺度上收集到的物候观测数据迅速累积,尽管已经在多个尺度上（物种、群落和景观尺度）观察到物候变化,但物候变化的机理仍然没有得到很好的理解。回顾了国内外植物物候研究的发展历程;总结了物候数据收集技术进展和全球物候变化的主要趋势;归纳了植物物候变化的机理与驱动因素;探讨了物候模型研究及物候对气候变化响应研究的主要方向。随着物候观测技术在不同尺度上应用的增加,物候研究进入了一个新的阶段。未来物候研究需要制定跨区域标准化观测指南,融合所有相关学科,改进物候模型,拓展研究区域;同时融合有效的历史物候资料,采用新技术和长期收集的物候数据为大数据时代植物物候学研究提供基础。     

全球变化下植物物候研究的关键问题

总结了全球变化下植物物候研究的主要进展,针对该领域国内外的几个热点问题进行了讨论。植物物候研究的重心从以前的野外观测和初步统计分析逐步过渡到以揭示物候周期的调控机制和环境效应为主,研究手段从植物物候对环境变化做出反应的表象描述转移到多尺度、多要素耦合关系的综合分析。随着学科交叉研究的不断深入,植物物候研究从植物个体及居群适应性研究转向植物物候变化对生态系统、气候演变、农业生产乃至人类健康等方面影响的系统评估。并且在该转变过程中出现了几个关键性问题,如不同温度带大气温度与光周期对植物物候期贡献力问题、植物物候变化对气候变暖的非线性响应特征、群落水平上植物物候研究的复杂性、以及农业生态系统中作物物候研究的重要性等。对我国植物物候研究现状和管理体系中亟待解决的问题提出了建议。     

植物物候对气候变化的响应

植物物候的变化可以直观地反映某些气候变化,尤其是气候变暖.植物生长节律的变化引起植物与环境关系的改变.生态系统的物质循环（如水和碳的循环）等过程将随物候而改变.不同种类植物物候对气候变化的响应的差异,会使植物间和动植物间的竞争与依赖关系也发生深刻的变化.目前欧洲、美洲、亚洲等许多地区均有关于春季植物物候提前,秋季物候推迟,使植物的生长季延长,从而提示气候变暖的趋势.植物物候的模拟模型构成生态系统生产力模型的重要部分.     

气候变暖对陆地植被-土壤生态系统的影响研究

由于自然因素及人类活动的长期影响,全球气候变化已经成为不容置疑的事实,并对陆地生态系统的植被及土壤产生了深远影响。陆地植被一土壤生态系统在全球气候变化中的反应与适应等过程已成为众多科学家所关注的问题。为更好地了解陆地植被一土壤生态系统对全球气候变化的响应机制,综述了气候变暖对植物的物候与生长、光合特征、生物量生产与分配,以及土壤呼吸等方面的影响,并对分析得到的结论进行了总结。分析指出,随着全球气候变暖,植物个体和群落特征以及土壤特性都会发生相应改变,高海拔地区的植被高度有增加趋势,而低海拔地区的植被可能出现矮化。然而,在以下方面还存有不确定性：（1）气候变暖导致的植被特征变化是否会减弱全球气候变化;（2）在较长时间尺度上气候变暖如何影响植物的物候和生长,特别是植物的体型;（3）高寒生态系统冬季土壤呼吸对气候变暖如何响应。     

甘南亚高山草甸群落的物候谱研究──兼论群落种多样性维持的机制

本文研究了亚高山草甸群落中常见的４２种植物的物候期。研究结果表明,此４２种植物在返青时间顺序、进入生殖生长的个体数量比例和进入生殖生长阶段的起始时间、历期上各不相同。根据返青顺序可将４２种植物分为三类。以单子叶禾莎草的返青时间为最早,多年生双子叶植物的返育时间较晚。根据进入生殖生长的个体占种群总个体数量的比例可分为：（１）生殖生长的个体比例在２０％以下,主要靠营养繁殖的种类;（２）具有双重繁殖特性的种类;（３）生殖个体占５０％以上的种类和（４）完全依靠有性繁殖的种类共四类。根据进入生殖生长的起始时间可分为三类。以上结果在一定程度上说明了植物种间由于在生长发育时间上的错位,减缓了种间对资源（如土壤矿物质营养）的竞争,使得群落中有多个种共存。     

干旱半干旱草地生态系统与土壤水分关系研究进展

研究干旱半干旱草地生态系统与土壤水分关系和相互作用机理对于揭示草地生态系统稳定性及其水土关键要素的变化过程具有重要意义。从不同界面、不同尺度综述了草地生态系统对土壤水分的影响及草地生态系统的响应与适应机制,总结了草地生态系统与土壤水分关系模型研究的相关进展,并分析了气候变化对草地生态系统和土壤水分关系的影响。草地生态系统通过影响水文过程和生态过程来影响土壤水分,土壤水分在植物生长发育、形态、生理生态过程、种间关系、群落组成和结构以及草地生态系统功能等方面对草地生态系统产生影响;充分揭示草地生态系统-土壤水分相互作用机理是模型研究的关键;气候变化对草地生态系统植物与土壤水分关系具有重要影响。今后应加强以下研究:1)开展草地不同优势种和植物功能型与土壤水分关系的研究,找出能反映植物对土壤水分响应的性状指标,阈值响应点及适应机制;2)注重对不同时间和空间尺度上的转换和比较;3)加强个体、群体和生态系统尺度草地植物生长模型的研究及其与土壤-植被-大气水分传输模型的耦合;4)加强草地生态系统与土壤水分关系对气候变化响应的研究。     

增温和氮素添加对内蒙古荒漠草原植物生殖物候的影响

植物物候是响应气候变化的敏感生物指示性状,在气候变暖并伴随着大气氮沉降增加的背景下,物候变化是气候变迁导致群落结构和生态系统功能变化的重要机制。本研究在以短花针茅(Stipa breviflora)为建群种的内蒙古荒漠草原设置了增温、氮素添加及其交互作用的实验平台,对短花针茅、木地肤(Kochia prostrata)、银灰旋花(Convolvulus ammannii)和细叶韭(Allium tenuissimum)4种多年生优势植物的开花、结实等物候期和生殖生长持续时间进行连续3个生长季(2008—2010年)的观察,植物物候观测采用定株标记,并对各植株上所有物候状态进行打分并以其非加权平均值作为植株整体物候分值,全年观测结束后将植株的物候分值随时间的变化利用Richards方程进行拟合,最终根据回归方程计算植物各物候期。结果表明:1)在3个生长季中,增温使4个植物种的平均开花、结实时间分别显著地提前了2.28和1.88 d·a-1;2)氮素添加对这些植物种的开花和结实时间都没有显著性影响;3)同时进行增温和氮素添加处理使植物的平均开花、结实时间分别提前1.75和1.53 d·a-1,但2种处理之间没有交互作用;4)增温使这些植物的平均生殖生长持续时间显著延长0.84 d·a-1,使短花针茅和木地肤生殖生长持续时间分别延长了1.20和3.25d·a-1。氮素添加使细叶葱的平均生殖持续时间显著缩短了1.52 d·a-1。这些结果揭示了气温升高和氮沉降影响下荒漠草原优势植物生殖物候的变化,将为分析不同种植物对各类资源需求规律的变化提供直接的实验证据,对于进一步预测全球变化背景下种间竞争格局和群落结构的演变趋势具有重要参考价值。     

基于OTCs模拟增温方式探讨气候变暖对青藏高原草地生态系统的影响

开顶式生长室(OTCs)增温实验是研究全球气候变化与陆地生态系统关系的主要方法之一,已广泛应用于青藏高原地区。该文通过对近些年国内外研究文献的回顾,分别从植物物候、群落结构、生物量和土壤方面综合分析青藏高原草地生态系统对OTCs模拟增温实验的响应。研究发现:增温使群落返青期提前、枯黄期延迟,生长季延长;有利于禾本科植物的生长;高寒草甸地下生物量分配格局向深层转移;高寒草地生态系统对模拟增温的响应存在不确定性,受到地域、群落类型和实验时间的影响;在增温条件下,降雨和冻土融化引起的土壤水分变化通过调控生态系统的物候、生产力、土壤等途径控制着生态系统对气候变暖的响应。并在此基础上,提出了将来应着重研究的几个方面。     

浙江古田山亚热带常绿阔叶林开花和结实物候的种间种内差异

植物物候学主要研究植物的生活史事件发生时间和环境因子之间的关系.物候作为植物重要的功能性状,却未见有在群落水平上将植物物候变异和群落结构特征(多度)联系起来探究植物物候变异规律的报道.为了探索物候性状的种间种内变异规律,2012年4月~2015年8月在中国东部地区浙江省古田山国家级自然保护区亚热带常绿阔叶林24hm~2大样地(GTS;29°10′19.4″~29°17′41.4″N,118°03′49.7″~118°11′12.2″E)内,对106种植物的物候进行连续观测,用标准差定性分析物候的种间和种内差异.结果表明,影响开花物候的气候因子为降水,群落开花高峰集中在5月,结实成熟高峰集中在10月.群落结实物候种间差异小于开花物候,其中结实物候种间差异为41天,开花物候种间差异为52天;群落水平植物开花和结实物候的种内变异小于种间;大部分物种开花物候的种内变异小于结实物候.本研究首次探讨了物候期种内变异系数和物种多度的关系,也是首次研究物候种内变异系数与物候期早晚的关系,发现群落物候种内变异与物种的多度不相关,群落开花物候种内变异与物候期的关系不显著,群落结实物候的种内变异和物候期具有显著负相关关系,即物候期早的结实物候种内变异大,物候期晚的结实物候种内变异小.将植物物候变异和群落结构特征(多度)联系起来探究植物物候变异规律,有利于理解物候的改变是怎样影响物种的相互作用和适合度,这对于在气候变化大背景下理解植物性状变异及物种分化和群落动态变化具有重要意义.     

北京地区气候变化和植被的关系——基于遥感数据和物候资料的分析

 气候变化对陆地生态系统的影响及其反馈是全球变化研究的焦点之一。本文利用1951～2000年的气温、降水等气候资料、1982～2000年的NOAA/AVHRR遥感数据和1951～2000年北京春季物候的代表性指标——山桃（Prunus davidiana）始花的物候数据,分析了在年际和年内时间尺度上北京地区各气候参量与植被变化之间的关系。结果显示：植物生长与温度之间的关系远比其与降水之间的关系密切;各气候参量和植被生长状况之间的关系因时间尺度而不同。1)月际水平上,具有显著生态学意义的气候指标对植被生长状况的影响更明显。2)温度与NDVI指标的相互作用最大为零时滞：年际水平上,影响时效约为1年;月际水平上,约为1个月。3)植物物候期与温度之间的关系远比其与降水之间的关系密切。年际尺度上,气候参量和植物物候期的相互作用是同时的,其中气温的影响时效为2年;月际尺度上,实际温度和植物物候期的相互作用时效约为1个月。     

Reproductive phenology of two co‐occurring Neotropical mountain grasslands

Aim

Climate tends to explain phenological variations in tropical ecosystems. However, water availability and nutrient content in soil strongly affect plant communities, especially those on old, climatically buffered, infertile landscapes (OCBILs), and may impact these ecosystems’ plant reproductive phenology over time. Here, we compare the reproductive phenology of sandy and stony tropical grasslands, two co‐occurring herbaceous communities of the campo rupestreOCBILs. We asked whether flowering, fruiting and dispersal are seasonal in both grasslands, and whether these phenophases differ due to variations in soil properties. We also asked whether the phenological strategies and the number of flowers and fruits differ between these two grasslands as soil conditions vary.

Location

Serra do Cipó, Minas Gerais, Brazil.

Methods

The phenology of herbaceous species of sandy and stony grasslands was monitored monthly over two consecutive years.

Results

Plants on sandy and stony grasslands flowered and fruited throughout the year. We did not find a distinct seasonal pattern at the community level of either studied grassland. However, flowering, fruiting and seed dissemination occurred in stony grasslands mainly during the rainy season, while sandy grassland species flowered in both seasons and fruited and disseminated seed mainly during the dry season, as observed in other savanna vegetation types in the Cerrado. Flower and fruit production was higher in sandy grasslands than in stony grasslands, which may be linked to higher water retention in sandy grassland soils. In both communities, species of Cyperaceae, Eriocaulaceae and Xyridaceae contributed most to overall production, whereas Poaceae and Velloziaceae, two important families in campo rupestre, barely participated in the reproductive phenology during our 2‐yr survey.

Conclusions

Despite a strong seasonal climate, there was no reproductive seasonal pattern at the community level in campo rupestre. This first investigation of Neotropical grassland phenology indicates that the differences in soil content may constrain the grassland reproductive phenology and restrict reproduction of stony grassland species to the most favourable season. Further studies of grassland phenology are necessary to disentangle the relative importance of soil, climate and other triggers, especially fire.     

Plant phenology and global climate change: Current progresses and challenges

Plant phenology, the annually recurring sequence of plant developmental stages, is important for plant functioning and ecosystem services and their biophysical and biogeochemical feedbacks to the climate system. Plant phenology depends on temperature, and the current rapid climate change has revived interest in understanding and modeling the responses of plant phenology to the warming trend and the consequences thereof for ecosystems. Here, we review recent progresses in plant phenology and its interactions with climate change. Focusing on the start (leaf unfolding) and end (leaf coloring) of plant growing seasons, we show that the recent rapid expansion in ground‐ and remote sensing‐ based phenology data acquisition has been highly beneficial and has supported major advances in plant phenology research. Studies using multiple data sources and methods generally agree on the trends of advanced leaf unfolding and delayed leaf coloring due to climate change, yet these trends appear to have decelerated or even reversed in recent years. Our understanding of the mechanisms underlying the plant phenology responses to climate warming is still limited. The interactions between multiple drivers complicate the modeling and prediction of plant phenology changes. Furthermore, changes in plant phenology have important implications for ecosystem carbon cycles and ecosystem feedbacks to climate, yet the quantification of such impacts remains challenging. We suggest that future studies should primarily focus on using new observation tools to improve the understanding of tropical plant phenology, on improving process‐based phenology modeling, and on the scaling of phenology from species to landscape‐level.     

Vegetation disturbance by fire affects plant reproductive phenology in a shrubland community in northwestern Patagonia, Argentina

Reproductive phenology is likely to vary spatially with environmental conditions that alter microclimate, in particular temperature. We hypothesized that within the same plant community type, environmental changes produced by recent burning would alter plant phenological patterns and temporal structure of the plant community. Specifically, we predicted accelerated flowering and fruiting dates in the burned, open environment compared with the unburned, intact community. We tested this hypothesis in a post-fire tall shrubland (matorral) in northwestern Patagonia, Argentina. During the reproductive season, phenological stages of seven vascular plant species were monitored weekly. Temperature, humidity, soil nutrients and photosynthetically-active radiation were also recorded. At the burned site, flowering began earlier in all species and the success rate of fruiting was higher. These patterns correlated with significant environmental differences, including higher mean temperatures at the burned site.     

Herbarium records identify sensitivity of flowering phenology of eucalypts to climate: Implications for species response to climate change

Flowering phenology is very sensitive to climate and with increasing global warming the flowering time of plants is shifting to earlier or later dates. Changes in flowering times may affect species reproductive success, associated phenological events, species synchrony, and community composition. Long‐term data on phenological events can provide key insights into the impacts of climate on phenology. For Australia, however, limited data availability restricts our ability to assess the impacts of climate change on plant phenology. To address this limitation other data sources must be explored such as the use of herbarium specimens to conduct studies on flowering phenology. This study uses herbarium specimens for investigating the flowering phenology of five dominant and commercially important Eucalyptus species of south‐eastern Australia and the consequences of climate variability and change on flowering phenology. Relative to precipitation and air humidity, mean temperature of the preceding 3 months was the most influential factor on the flowering time for all species. In response to a temperature increment of 1°C, a shift in the timing of flowering of 14.1–14.9 days was predicted for E. microcarpa and E. tricarpa while delays in flowering of 11.3–15.5 days were found for E. obliqua, E. radiata and E. polyanthemos. Eucalyptus polyanthemos exhibited the greatest sensitivity to climatic variables. The study demonstrates that herbarium data can be used to detect climatic signals on flowering phenology for species with a long flowering duration, such as eucalypts. The robust relationship identified between temperature and flowering phenology indicates that shifts in flowering times will occur under predicted climate change which may affect reproductive success, fitness, plant communities and ecosystems.     

Plant species with the trait of continuous flowering do not hold core roles in a Neotropical lowland plant‐pollinating insect network

Plant–animal interaction science repeatedly finds that plant species differ by orders of magnitude in the number of interactions they support. The identification of plant species that play key structural roles in plant–animal networks is a global conservation priority; however, in hyperdiverse systems such as tropical forests, empirical datasets are scarce. Plant species with longer reproductive seasons are posited to support more interactions compared to plant species with shorter reproductive seasons but this hypothesis has not been evaluated for plant species with the longest reproductive season possible at the individual plant level, the continuous reproductive phenology. Resource predictability is also associated with promoting specialization, and therefore, continuous reproduction may instead favor specialist interactions. Here, we use quantitative pollinating insect–plant networks constructed from countryside habitat of the Tropical Wet forest Life Zone and modularity analysis to test whether plant species that share the trait of continuous flowering hold core roles in mutualistic networks. With a few exceptions, most plant species sampled within our network were assigned to the role of peripheral. All but one network had significantly high modularity scores and each continuous flowering plant species was in a different module. Our work reveals that the continuous flowering plant species differed in some networks in their topological role, and that more evidence was found for the phenology to support specialized subsets of interactions. Our findings suggest that the conservation of Neotropical pollinating insect communities may require planting species from each module rather than identifying and conserving network hubs.     

Benefits of hyperspectral remote sensing for tracking plant invasions

Aim We aim to report what hyperspectral remote sensing can offer for invasion ecologists and review recent progress made in plant invasion research using hyperspectral remote sensing. Location United States. Methods We review the utility of hyperspectral remote sensing for detecting, mapping and predicting the spatial spread of invasive species. We cover a range of topics including the trade‐off between spatial and spectral resolutions and classification accuracy, the benefits of using time series to incorporate phenology in mapping species distribution, the potential of biochemical and physiological properties in hyperspectral spectral reflectance for tracking ecosystem changes caused by invasions, and the capacity of hyperspectral data as a valuable input for quantitative models developed for assessing the future spread of invasive species. Results Hyperspectral remote sensing holds great promise for invasion research. Spectral information provided by hyperspectral sensors can detect invaders at the species level across a range of community and ecosystem types. Furthermore, hyperspectral data can be used to assess habitat suitability and model the future spread of invasive species, thus providing timely information for invasion risk analysis. Main conclusions Our review suggests that hyperspectral remote sensing can effectively provide a baseline of invasive species distributions for future monitoring and control efforts. Furthermore, information on the spatial distribution of invasive species can help land managers to make long‐term constructive conservation plans for protecting and maintaining natural ecosystems.     

Linking flowering and reproductive allocation in response to nitrogen addition in an alpine meadow

Aims Plants can change in phenology and biomass allocation in response to environmental change. It has been demonstrated that nitrogen is the most limiting resource for plants in many terrestrial ecosystems. Previous studies have usually focused on either flowering phenology or biomass allocation of plants in response to nitrogen addition; however, attempts to link flowering phenology and biomass allocation are still rare. In this study, we tested the effects of nitrogen addition on both flowering phenology and reproductive allocation in 34 common species. We also examined the potential linkage between flowering time and reproductive allocation in response to nitrogen addition.Methods We conducted a 3-year nitrogen addition experiment in Tibetan alpine meadow. We measured first flowering date and the reproductive allocation for 34 common plant species in control, low and high nitrogen added plots, respectively. One-way analysis of variance was used to examine differences of first flowering date and reproductive allocation among treatments. The relationships between the change in species first flowering date and change in reproductive allocation in response to nitrogen addition were examined by calculating Pearson correlation coefficients.Important findings For most species, both first flowering date and reproductive allocation significantly responded to nitrogen addition. Nitrogen addition significantly delayed the first flowering date and reduced the reproductive allocation for all graminoid species, but accelerated flowering and increased reproductive allocation for most forb species. We found that changes in first flowering date significantly negatively correlated with the changes in reproductive allocation over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. Our results suggest that species-specific switch from vegetative growth to reproductive growth could influence species performance.     

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3‐year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow‐rooted, early‐flowering plant, which may be mainly constrained by upper‐soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.